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Life-Histories of the Frogs of
Okefinokee Swamps, Georgia

7^

Life-Histories of the Frogs of
Okefinokee Swamp, Georgia

NORTH AMERICAN SALIENTIA

(ANURA) NO. 2

By

Albert Hazen Wright

New York
The Macmillan Company

1931

COPYRIGHT, 1032

BY ALBERT HAZEX WRIGHT

prixted by
The Caycga Press

TO DR. LEONHARD STEJNEGER

NESTOR OF AMERICAN HERPETOLOGY

AND KINDLY COUNCILLOR OF

YOUNGER BIOLOGISTS

Field Work Under Heckscher Grants Nos. 14 and 31

FOREWORD

In 1 9 14 we wrote as an introduction to a similar work for Ithaca, N. Y.,
the following paragraphs :

"Notwithstanding the extensive use of our American Anura for anatomical
and embryological purposes, comparatively little attention has been given to
the collection of definite data bearing upon the breeding habits and larvae of
this group. In this connection, Boulenger (1897), in the preface to his work
upon 'The Tailless Batrachians of Europe', observes:

*I would also express a hope that a Uttle book of this kind . . . may
have the effect of stimulating interest in a subject that has been too much
neglected, and in the cultivation of which new workers will find much to repay
their efforts, especially if applied in other regions of the globe, which, though
richer in Batrachians, have as yet yielded Httle or nothing to our knowledge
of the life-histories.'

"Dr. Gill (1898), in his review of the work just mentioned, remarks that
he hopes this monograph* 'may serve as a model for other lands, and not
least for the United States. . . . Every sojourner in the country must have
noticed the masses of transparent jelly-like spheres in the water, but none in
the United States could refer such masses with certainty to the parent species.'
To a local study of these very phases of Anuran hfe, my investigations for the
past seven or eight years have been directed."

Since 1914 we have continued these studies from Ithaca to California.
With these studies of the southeast, Storer's excellent accounts of the Pacific
coast forms and Strecker's long continued notes on Texas species, the Hfe
histories of most of the Salientia of North America are now outlined. Some
remain untouched and many details need to be filled in the outhnes at hand.

*The material upon which this report is based is from two sources: the personal material
collected in Okefinokee Swamp from 1909-1922; and the United States National Museum
collection of the Okefinokee species herein considered. Before the completion of this manu-
script Mrs. Wright and I examined also all of the U. S. National Museum frogs of North
America north of the Mexican line. This material furnished comparative notes on the
following topics: the range; structural differences between sexes; least and greatest breeding
sizes; transformation data and sizes; measurements of frogs from transformation to largest
adults for rate of growth; first appearance and autumnal disappearance dates; and other
incidental data. Additional information came from the U. S. National Museum's un-
identified and uncatalogued tadpoles and transformation stages which the present author
is examining from time to time. To Dr. Leonhard Stejneger, Dr. Remington Kellogg, and
Miss Doris Cochran we owe countless kindnesses.

To Doubleday, Doran & Co. we are indebted for permission to use excerpts from Miss
M. C. Dickerson's useful work, The Frog Book.

The investigation upon which this work is based was supported in 1921 and 1922 by a
grant from the Heckscher Foundation for the Advancement of Research, established at
Cornell University by August Heckscher.

CONTENTS

PAGE

Foreword v

Table of contents vii

List of illustrations xi

Introduction i

Acknowledgments i

To residents i

To specialists 3

Previous pertinent amphibian explorations toi9i4 3

Subsequent explorations (i9i4toi92 2) 9

The Okefinokee region 10

Plant habitats 11

A catalogue of the species of Salientia of the Okefinokee region (1922) . . 13

Hypothetical species 15

General discussion of the Okefinokee Salientia 15

Range 15

Coloration of spirit specimens (1912). Measurements (1912-1914) . . 16
Measurements of Okefinokee frogs at 28, 44, 66, 82, 95, 113, 125, 136,

150, 180 mm 16

First appearance and autumnal disappearance 24

Breeding sizes 24

Voice 26

Mating 26

Coloration from life 27

Structural differences 28

Customary amplexation 28

Cross embraces 28

Departures from customary amplexation and their significance ... 29

Ovulation 30

The eggs 33

Synopsis of the eggs of the Okefinokee Salientia 35

Egg-laying process 37

Hatching period 37

Larval period 38

Tadpoles 39

Synopsis of the tadpoles of the Okefinokee region 41

Transformation period 56

Transformation size 58

Rate of growth 59

Food 62

Enemies 62

Affinities ' 62

Criteria i-ii 63

Hypotheses 65

Summary of affinities 66

Suggestions for local or resident naturalists and collectors in the south-
eastern United States 67

General topics 67

Special problems for each species 68

vu

3933r>

viii Frogs of the Okefinokee Swamp

PAGE

General accounts of species 73

Life-history of the Spade-foot 5cap/(iopMs/ioZ6rooH{ (Harlan) ... 73

Scientific name 73

Common names 73

Range . 73

Checklist — Stejneger and Barbour 1923 73

Supplementary records 73

Local Okefinokee records 74

General appearance ■ 75

Measurements 75

Habitat 76

First appearance 76

General habits 77

Voice 86

Mating 90

Coloration of male from life . 90

Coloration of female from life 90

Structural differences 90

Duration, day or night 91

Amplexation (normal, abnormal cross) 92

Ovulation 92

Habitat 92

Period 92

Temperature and humidity 93

Egg-laying process 93

Eggs 95

Attachment 95

Egg mass 95

Egg description 95

Dangers 97

Hatching period 98

Mature tadpole 98

Color description from life 98

General appearance 98

Mouth parts 99

Measurements 99

General remarks 99

Larval period loi

Transformation 102

Period 102

Size 102

Growth 103

Food 103

Autumnal disappearance 104

Affinities 105

Bibliography 106

Life-history of the oak toad {Bufo quercicus Holbrook 107

This species and subsequent accounts have the topics of the spade-foot
with these added topics

Coloration of spirit specimens (191 2) 109

Structural characters (191 2) no

Measurements (191 2) m

Contents ix

PAGE

General habits 114

Metachrosis 114

Variations in color 114

Life-history of the southern toad Bufo terrestris Bonnaterre 133

Life-history of the cricket-frog i4ms ^ryZZws (Le Conte) 156

Life-history of the black chorus frog. Pseudacris nigrita (Le Conte) . 196
Life-history of the smooth chorus frog. Pseudacris occidentalis (Baird and

Girard 204

Life-history of the little chorus frog. Pseudacris ocularis (Holbrook) . . 206

Life-history of the ornate chorus frog. Pseudacris ornata (Holbrook) 227

Life-history of the Anderson tree-frog. Hyla aridersonii Baird . . . . 227

Life-history of the green tree-frog. Hyla cinerea (Schneider) .... 248

Life-history of the pine wood's tree-frog. Hyla femoralis Latreille. . . 272

Life-history of the Florida tree-frog. Hyla gratiosa Le Conte 294

Life-history of the southern tree-frog. Hyla squirella Latreille 310

Life-history of the common tree-toad. Hyla versicolor (Le Conte). . . 325

Life-history of the gopher frog. Rana aeso-pus (Cope) 334

Life-history of the green frog. Rana clamitans Latreille 352

Life-history of the southern bull frog. Rana grylio Stejneger 364

Life-history of the Heckscher's frog. Rana heckscheri Wright 384

Life-history of the mink frog. Rana septentrionalis Baird 397

Life-history of the southern leopard frog. Rana sphenocephala {Co-pe) . . . 419

Life-history of the carpenter frog. Rana virgatipes Cope 435

Life-history of the narrow-mouthed toad. Gastrophryne carolinensis

(Holbrook) 452

LIST OF ILLUSTRATIONS

(Plates mounted by A. A. Wright)

1 92 1 photographs by F. Harper and the author.

1922 photographs by F. Harper and A. A. Wright.
1923-29 photographs by A. A. and A. H. Wright.

(Photographs developed and printed by Troy Studio
Engravings by The Hurst Engraving Co.)

Plate I. Lateral aspects. Bufo, Gasirophryne, Pseudacris, Scaphiopus. 14

1. Bufo americanus. 6. Gastrophryne carolinensis.

2. Bufo fowleri. 7. Pseudacris ocularis.

3. Bufo terrestris. 8. Pseudacris ornala.

4. Scaphiopus holbrookii. 9. Pseudacris nigrita.

5. Bufo quercicus.

Plate II. Lateral aspects.

1. Acris gryllus.

2. H])la crucifer.

3. Hyla andersonii.

4. Hyla gratiosa.

5. Hyla cinerea.

Hyla and Acris

6. Hyla evittata.

7. Hyla squirella.

8. Hyla femoralis.

9. Hyla versicolor.
10. Hyla avivoca.

Plate III. Lateral aspects.

1. Rana septentrionalis.

2. Rana clamitans.

3. Rana virgatipes.

4. Rana heckscheri.

Rana males

5. Rana aesopus.

Rana pipiens.
Rana sphenocephala.
Rana grylio.

Plate IV. Amplexation

Normal types.

1. Pectoral. Ranidae. Rana pipiens.

2. Inguinal. Scaphiopodidae. Scaphiopus holbrookii.

3. Axillary. Hylidae. Hyla versicolor.

4. Axillary. Brevicipitidae. Gastrophryne carolinensis.

30

Abnormal types.

5. Preaxillary. Gastrophryne carolinensis.

6. Multiple amplexation. Pseudacris ocularis.

7. Cross-embrace. Bufo valliceps and Rana pipiens.

8. Two males in embrace. Hyla versicolor.

Plate V. Normal embraces.

30

1. Hyla cinerea. Axillary. 7.

2. Hyla femoralis. Axillary. 8.

3. Hyla gratiosa. Axillary. 9.

4. Hyla squirella. Axillary. 10.

5. Acris gryllus. Axillary. 11.

6. Pseudacris ocidaris. Axillary. 12.

Bufo quercicus. Axillary,
Bufo terrestris. Axillary.
Gastrophrijne carolinensis. Axillary.
Bufo fowleri. Axillary.
Rana clamitans. P^ectoral.
Rana sphenocephala. Pectoral.

Plate VI. Egg masses.

1. Acris gryllus.

2. Scaphiopus holbrookii.

Gastrophryne carolinensis.
Pseudacris ocidaris.

30

xii Frogs of the Okefinokee Swamp

Plate VII. Eggs of Hylas 38

1. Hyla cinerea. 4. Hyla cinerea.

2. Hyla versicolor. 5. Hyla squirella.

3. Hyla femoralis. 6. Hyla gratiosa.

Plate VIII. Egg masses 38

1. Rana clamitans. 4. Rana grylio.

2. Bufo quercicus. 5. Biifo terrestris.

3. Bufo quercicus. 6. Bufo terrestris.

Plate IX. Rana egg masses 38

1. Ra7ia septentrionalis. 3. Rana sphenocephala.

2. Rana virgatipes. 4. Rana aesopus.

Plate X. Mature tadpoles 46

1. Hyla andersonii. 8. Pseudacris ocularis.

2. Hyla cinerea. g. Hyla versicolor.

3. Hyla crucifer. 10. Hyla versicolor.

4. Hyla femoralis. 11. Acris gryllus.

5. Hyla gratiosa. 12. Scaphiopus holbrookii.

6. H^Jla gratiosa. 13. Bufo quercicus.

7. Hyla squirella. 14. Gastrophryne carolinensis.

Plate XI. Mature tadpoles. {Rana.) 46

1. Rana aesopus. 8. Rana heckscheri.

2. Rana aesopus. 9. Rana sphenocephala.

3. Rana clamitans. 10. Rana sphenocephala.

4. Rana clamitans. 11. Rana septentrionalis.

5. Rana grylio. 12. Rana pipiens.

6. i^ana grylio. 13. iiana virgatipes.

7. /^ona heckscheri. 14. /2ana palustris.

Plate XII. Tadpole mouthparts. {Hyla, Acris, Pseudacris, Bufo) . 54

1. i/2/Za crucifer. 8. Pseudacris ocularis.

2. //i/Za gratiosa. 9. /f^/Za squirella.

3. /^2/Za andersonii. 10. 5m/o americanus.

4. Hyla cinerea. 11. Bufo terrestris.

5. Pseudacris feriarum. 12. Hyla versicolor.

6. Acris gryllus. 13. Hyla femoralis.

7. Bufo fowleri.

Plate XIII. Tadpole mouthparts. {Rana) 54

1. i^awa caiesbeiana. 4. i^awa aesopus.

2. flana sphenocephala. 5. i^ana pipiens.

3. i2ona palustris.

Plate XIV. Tadpole mouthparts {Rana, Gastrophryne, Scaphiopus). . 54

1. jRana heckscheri. 5. 72ana grylio.

2. Foun^ i^ana heckscheri. 6. Gastrophryne carolinensis.

3. i^ano septentrionalis. 7. i?ana clamitans.

4. /2awa virgatipes. 8. Scaphiopus holbrookii.

Plate XV. Transformed frogs 62

1. /fj/^^ andersonii. 8. Pseudacris ocularis.

2. Hyla cinerea. 9. Scaphiopus holbrookii.

List of Illustrations xiii

Hyla femoralis. lo. Gastrophrtjne carolinensis.

Hyla gratiosa. ii. Bufo terrestris.

Hyla sqtdrella. 12. Bufo quercicus.

Hyla versicolor. 13-16. Typical series, tadpole to trans-
oms gryllus. formed frog. Hyla gratiosa.

Plate XVI. Transformed frogs (Ranas) 62

1. Rana virgatipes. 6. Rana grylio.

2. Rana aesopus. 7. Rana sphenocephala.

3. Rana grylio. 8. Rana sphenocephala.

4. Rana clamitans. g. Rana septentrionalis.

5. Rana clamitans. 10. Rana septentrionalis.

Plate XVII. Chart of growth 62

Transformation sizes in comparison with greatest mature sizes of males and
females of twenty species.

Plate XVIII. Spade-foot {Scaphiopus holbrookii) 78

1. Male and female. Dorsal. 4. Male. Lateral.

2. Male and female. Ventral. 5. Egg masses.

3. Adult. Lateral. 6. Eggs, stained.

Plate XIX. Spade-foot {Scaphiopus holbrookii) 94

I. Habitat. 6.7. Tadpoles.

2.3.4. Males croaking. 8.9.10.11. Tadpoles and transforming

5. Eggs. frogs.

Plate XX. Oak toaj) {Bufo quercicus) no

I. Habitat. 6. On sand.

2-5. Males croaking. 7. Egg string.

Plate XXI. Oak toad {Biifo quercicus) 126

I. Habitat. 6. Egg bars, stained.

2.3. Toads in daytime retreats. 7.8.9. Tadpoles.

4. Female and male. Ventral. 10. Transformed toad.

5. Clusters of egg bars.

Plate XXII. Southern toad {Bufo terrestris) 142

1. Habitat. 8.9. Tadpoles.

2. Female and male. Ventral. lo.ii. Transformed toads.

3.4.5. Males croaking. 12. Egg strings, stained.

6. Male and two females. Lateral.

7. Biifo quercicus and Btifo terrestris compared.

Plate XXIII. Cricket-frog {Acris gryllus) 158

I. Habitat. Chase Prairie. 8.9.10. Males croaking.

2.3. Tadpoles. 11. Eggs.

4. Tadpoles with four legs. 12. Tadpoles with two legs.

5.6. Transforming and transformed frogs.

7. Two males and two females. Ventral.

Plate XXIV. Black chorus frog {Pseudacris ocularis). .... 206

I. Habitat. Saddlebag Pond. 8. Tadpole with short tail.

2.3. Males, croaking. 9. Transformed frog.

4.5 Mature tadpoles. 10. Male croaking.

6. Tadpole with two legs. 11. Habitat, near Siren Run.

7. Tadpole with four legs. 12. Male croaking.

xiv Frogs of the Okefinokee Swamp

Plate XXV. Anderson's tree-frog ( //y?a andersowu) 238

1. Habitat. Lakehurst, N. J. 4. Tadpole.

2. Male. Ventral. 5.6.7.8.9. Series, tadpole to trans-

3. Male. Lateral. formed frog.

Plate XXVI. Green tree-frog ( /fi/ia cmerea) 254

I. Habitat. Billy's Lake. 6. Male and female. Ventral.

2.3. Males, croaking. 7. Small tadpole.

4. Male and female. Lateral. 8. Mature tadpole.

5. Male, croaking.

Plate XXVIL Pine woods tree-frog (Hylafemoralis) 270

1. Male croaking. 6. Transformed frog.

2. Male resting on bit of bank. 7. Eggs, stained.

3. Male croaking. 8. Female and male.

4. Tadpole. 9- Eggs, stained.

5. Two males. Lateral.

Plate XXVIIL Pine woods tree-frog {Hylafemoralis) 286

1.2. Pools with egg-clusters. 9. Tadpoles with four legs.

3. Eggs. 10. Tadpole with shrinking tail.

4.5. Males, croaking. 11. Tadpole with stub of tail.

6.7.8. Tadpoles. 12. Transformed frog.

Plate XXIX. Florida tree-frog ( i/?/?a ^mhosa) 294

1. Habitat. 6. Mature tadpole, (abnormal).

2. Frog on sand. 7. Mature tadpole, (normal).
3.4.5. Males, croaking.

Plate XXX. Florida tree-frog ( //yZa ^mhosa) 302

1. Coat-bet pond. 6. Pair on log.

2. Male, semi-inflation. Lateral. 7. Male.

3.4. Males, showing effect of rubbing. 8. Egg mass in a pan.
5. Female and male.

Plate XXXI. Squirrel tree-frog ( i/?/?o sgwiVeZZa) 310

1.2. Males, croaking. 12. Half-grown frog. Lateral.

3-9. Tadpole series. 13. Male. Ventral,

lo.ii. Transformed frogs.

Plate XXXII. Squirrel tree-frog ( /f?/Za sgmVeZ/a) 318

1. Anna's pond. 4-5-6. Males, croaking.

2. Mated pair. 7. Female and male.

3. Egg mass. 8. Eggs.

Plate XXXIII. Trei:-to ad (Hyla versicolor) 326

1. Old turpentine pit at Camp 4. Male, croaking.
Pinckney. 5. Transformed frog.

2. Male, croaking. 6. Egg packet.

3. Male. 7- Female and male.

Plate XXXIV. Gopher frog {Raiia aesopus) 334

1. Cypress pond. 4- Female. Ventral.

2. Male, croaking. 5. Frog at entrance of gopher-turtle

3. Male. Ventral. hole.

6. Eggs, stained.

List of Illustrations xv

Plate XXXV. Gopher frog {Rana aesopus) 350

1. Gopher-turtle hole. 3. Male on sand.

2. Male at base of pine. 4-6. Tadpole series.

Plate XXXVI. Greh:^ fuog {Rana clainitans) 358

1. Egg mass. 6. Adult female and transformed frog.

2. Eggs, stained. 7.8. Transformed R. clamitans and
3.4. Tadpoles. R. grylio compared.

5. Transformed frog. 9. Male, croaking.

10. Male. Ventral.

Plate XXXVII. Southern bullfrog (Rana grylio) 366

1. Male. Ventral. 5.6. Tadpoles.

2. Female. Ventral. 7. Transformed frog.

3. Transforming frog. 8. Site of egg mass.

4. Male, croaking. 9. Bullfrog pond.

Plate XXXVIII. River-swamp frog (Rana heckscheri) 382

I. Thompson's Landing. 4.5. Mature frog. Lateral.

2.3. Mature tadpoles. 6. Mature frog. Ventral.

Plate XXXIX. Mink frog (Rana septentrionalis) 398

1. Male. Lateral. 4- Egg mass.

2. Female. Ventral. 5. Transforming frog.

3. Hellgate pond, Onekio, N. Y. 6. Adult.

Plate XL. Southern leopard frog (i^ana spAenocepAaZo) 422

1. Male, croaking. 5. Female. Ventral.

2. Eggs, stained. 6. Female. Lateral.

3. Male, croaking. 7. Male. Lateral.

4. Egg mass in camp.

Plate XLI. Southern leopard frog (Rana sphenocephala) 430

I. Siren Run, Billy's Id. 6. Tadpole with stub of tail.

2.3.4.5. Tadpoles. 7. Transformed frog.

Plate XLII. Carpenter frog (Rana virgatipes) 438

1. Second "Dreen," Chesser Id. 4-S-6. Males croaking.

2. Egg mass. 7. Male. Lateral.

3. Eggs, stained. 8. Female, Lateral.

Plate XLIII. Carpenter frog (Rana virgatipes) 446

1. Male. Ventral. 4. R. grylio transformed and R.

2. Female. Ventral. virgatipes compared.

3. Eggs in situ. 5. Adult. Ventral.

6.7. Tadpoles.

Plate XLIV. Narrow-mouthed toad (Gastrophryne carolinensis) . . . 462

1. Pool, Billy's Id. 5.6.7. Tadpoles.

2. Male and Female. Lateral. 8. Transformed frog.

3. Female and male. Ventral. 9. 10. 11. Males, croaking.

4. Egg-clusters. 12. Eggs, stained.

Plate XLV. Map of Okefinokee Swamp, Ga 478

TEXT FIGURES
I. Individual eggs.

INTRODUCTION

In 1915 we wrote: "The object of all the Cornell (1909-1914) expeditions
was to study and put on record something of the biological conditions in this
extensive fresh-water swamp, which still presents in a large measure primitive
and interesting conditions of environment, before they should become forever
changed by the now rapidly penetrating lumbermen."

From these private trips we published A Biological Reconnaissance of the
Okefinokee Swamp in Georgia. (The Birds by A. H. Wright and Francis
Harper, 1913 ; The Turtles, Lizards and AlHgators by A. H. Wright and W. D.
Funkhouser, 191 5; The Snakes bj^ A, H. Wright and S. C. Bishop 1915; The
Fishes, by E. L. Palmer and A, H. Wright, 1922). Besides these, three more
reports were prepared but not published, namely. The History of the Swamp
by A. H. Wright, The Amphibia by A. H. Wright and Julia Moesel, and The
Reptiles (2nd Report) by S. C. Bishop and A. H. Wright.

In 1 92 1 the author was given a grant from the Heckscher Foundation for
the Advancement of Research established at Cornell University and this was
continued for the season of 1922 (see topic Subsequent Explorations, 1914
to 1922). This paper therefore includes notes and material from 1909-1922, —
the period to 1914 or later in the center of the swamp before lumbering; the
period 192 1 with lumbering (Billy's Island the base); and the period 1922 on
Chesser Island to have conditions more Hke those of 19 14 and to study the
surrounding biota of the swamp.

ACKNOWLEDGMENTS TO RESIDENTS
In 191 5 we held that most valuable services were rendered to this party
by the Lee family, (Mrs. Lee, the mother of the first generation, Jackson,
David, Bryant, Farley, Harrison, Joseph of the second generation, and
Marion and Ely of the third generation), living on Billy's Island, the only
human inhabitants of the interior of the swamp, and practically out of touch
with the outside world. Their primitive mode of living had adapted them to a
marvelous degree to the appreciation of the wild life about them, and their
observations and knowledge of natural phenomena proved to be surprisingly
accurate. The older men and boys were indispensable as guides while the
party was in the swamp, and on the exit of the party a container was left
with them to be filled with specimens which might come to their hands later in
the year. This container, full of material chosen with evident care and good
judgment, was received November 15, 19 12, and the specimens thus secured
proved a valuable addition to those previously collected. In December, 19 13,
data were obtained on the winter condition of some of the forms here noted.
In 1 92 1 Jackson, Harrison and David Lee were in the swamp and rendered in-
valuable assistance. From the next generation we also received continued service.
These were Henry Harrison and Alarion Lee and their brothers and cousins.

2 Frogs of the Ohefinokee Swamp

In 192 1 we entered the swamp with letters of introduction from the
Hebards of the Hebard Cypress Company. No one received finer courtesies
or consideration than we all the while we were in their midst. We camped in
the lumber settlement of 300 or 400 or more. The superintendents (John M.
Hopkins, A. J. Armstrong), gave instructions and courtesies which lightened
in many ways our search. To the superintendents on the island (H. S.
Quarterman and V. Alex, Quarterman), to the boarding house (Mr. Wm, J.
Wilson and his sympathetic women helpers), to the timber cutters, skidder
force, pile driver force, girdling crew, commissary or store unit, the shop
employees, we extended our thanks for countless favors. To eat and associate
with this roster of real workers who were truly tolerant of frogcatchers, snake
"doctors" and other such queer researchers as we is an opportunity and
experience we will never have again. As irregular as we might be in our
eating hours, Mrs. Wilson and her force were patient. To one group we owe
much help. They are the train force. These twelve or fifteen men (engi-
neers, firemen, flagmen) would stop a train in impossible places to pick us
up weary and often laden, or with engines in rare cases come for us at distant
points. For no one on Billy's Island's roster did the author feel more in-
debted than to Sam Edwards, the time keeper and in many ways chief
factotum. He was a well read man who could well understand the import of
our mission. All in interest, fine spirit, assistance and information contributed
to the attainment of our material.

This official roster deserves considerable thanks but rest assured that we
were wise enough to enlist as well the interest of the Quarterman, Edwards,
Strickland, Woodward, Jordan, Craven, Carter, Cox, Bennett and other
Juniors who were volunteer assistants.

Continually we interviewed the older inhabitants whose intimate ex-
periences with the fife of the region proved invaluable. Each of the Lee
brothers and the next generation as well we thus quizzed from the mammals
to the fishes. In the same way R. A. Chesser and his brother Sam with their
sons Ben, Harry, Ridley and Tom have unsparingly helped and wiUingly
recounted their knowledge and experiences.

Messrs. James Henderson, Walter Davis, J. D. Hendrix, Julius Godwin,
Norman Godwin, each contributed notes as we went down the catalogue of
forms from fish to mammals and many a long session ensued.

Two members of the Hebard Company who particularly had rich knowl-
edge of the swamp, its out of way parts and its rare inhabitants were par-
ticularly Sam Mizzell and his brother Hamp Mizzell.

In 1922 we were the guests of the Chessers on Chesser Island and to
them we can never fully express the joyous and profitable summer they gave
us. In 1922 we returned to Billy's Island for a week or more and received
countless favors from our old friends of 1921 or of 1912-1921. All in all the
animal experiences related to us by the inhabitants and members of the
lumber colony as well were very valuable, particularly so for the birds and
mammals. For the cold blooded animals they had less observations. Never-
theless they knew where most of them occurred and helped in securing them.

Introduction 3

To several people of Folkston and Wayeross we owe kindnesses, particularly
to Mrs. J. L. Walker and her daughter of Wayeross, Georgia.

ACKNOWLEDGMENTS TO SPECIALISTS

To the constant interest and assistance of Mrs. Anna Allen Wright, my
wife, attaches much of whatever merit this paper possesses. She was on the
trip in 1922. In 19 12 she helped to carry part of my teaching in nvy ab-
sence, and in 192 1 purposed to be my associate. She has made the drawings,
mounted the plates and helped Dr. Francis Harper in the photography in 1922.

Both in 192 1 and in 1922 Dr. Francis Harper was on the trip. He gave
intelligent interest in collecting material and notes and was my chief reUance
in the photography of the frogs in 192 1. In 1922 Dr. Miles D. Pirnie assisted
me in general, Mrs. Wright in plant collecting and identifications, and Dr.
Harper in the preparation of skins.

At the American Museum of Natural History the particular species sought
on two different occasions were kindly placed at our disposal by Dr. G.
Kingsley Noble, Messrs. C. D. Pope and Wm. Hassler. At the Museum of
Comparative Zoology, Dr. T. Barbour and Mr. A. Loveridge made my search
very expeditious and profitable. In the same way, several days spent on
material at the University of Michigan proved pleasant and of interest,
thanks to Dr. A. G. Ruthven, and Dr. F. M. Gaige, but particularly to Mrs.
Helen Thompson Gaige. To M. Graham Netting of Carnegie Museum, Pitts-
burg, Pa. also goes our indebtedness for favors deeply appreciated.

PREVIOUS PERTINENT AMPHIBIAN EXPLORATIONS TO 1914

In 1 9 13 we wrote the following general introduction to a contemplated
paper on 'The Biological Reconnaissance of the Okefinokee Swamp in Geor-
gia. The Amphibians. By A. H. Wright and Juha Moesel." It never was
pubhshed but excerpts from it appear throughout this work. In it the fol-
lowing introduction appeared.

For several decades the state of Georgia has received very little herpe-
tological attention yet the history of North American batrachology proves it
to be one of the most important in the Union. The most recent pertinent
hst comes from the immediate southeast in Florida. In 1911-1912, Mr.
Richard F. Deckert collected the following Salientia near Jacksonville.
They are'

*Ranapipiens *Hyla pickeringii

Rana sphenocephala *Hyla squirella

*Rana aesopus Hyla femorahs

Rana clamitans Hyla cinerea

Rana catesbiana Hyla gratiosa

Rana grylio *Chorophilus nigritus

Engystoma carolinense *Chorophilus occidentahs

Bufo lentiginosus Chorophilus ocularis

Bufo quercicus Acris gryllus

' Deckert, R. F. Copeia No. 3, Feb. 19, 1914; No. 5, April 15, 1914; No. 9, Aug. 29, 1914.

4 Frogs of the Okefinokee Swamp

At an earlier date and slightly farther south from Green Cove Springs and
Hastings, Fla., C. S. Brimley lists-

Plethodon glutinosus *Chorophilus ornatus

*Desmognathiis fusca auriculata Hyla cinerea

Diemyctylus viridescens *Hyla squirella

Amphiuma means Hyla gratiosa

Siren lacertina Hyla femorahs

Bufo quercicus *Rana aesopus

Acris gryllus *Rana eatesbiana

At Nashville and Allapaha, Ga. to the northwest of the Okefinokee
Swamp, Mr. W, J. Taylor secured the following forms:'

Bufo lentiginosus lentiginosus Rana virescens sphenocephela

Acris gryllus gryllus *Rana virescens virescens

Chorophilus Occident alis *Rana areolata areolata

Hyla squirella Rana clamata

Hyla femoralis *Desmognathus fusca

Hyla carolinensis *Amphiuma means

Thus, from these three lists we could compile a hypothetical (*) list of
forms to be sought in Okefinokee Swamp or its environs in the year 1913.
The following 1 1 forms might be thought the nucleus of it. They are

Rana pipiens Chorophilus nigritus

Rana areolata areolata Chorophilus occidentalis

Rana aesopus Chorophilus ornatus

Rana catesbeiana Desmognathus fusca

Hyla squirella Desmognathus fusca auriculata

Hyla pickeringii

In the earlj-- part of the last century no state of the United States fur-
nished more amphibian material than Georgia and its earUer herpetological
writers and coUecters included such worthies as Major J. LeConte, Dr. J. L.
LeConte, Dr. Holbrook and his east Georgian correspondents and collectors
like Dr. Couper of St. Simon's Island and Dr. Harden of Riceboro, Ga. This
latter place, the type locality of five or six mammals is equally interesting
in batrachology. Here the LeConte plantation is situated and no doubt it is
the type locality for some of the Georgian species Maj. J. LeConte described.

In 1825 (May 16) he distinctly records three forms from Georgia^:

Hyla lateraHs Hyla femorahs

Hyla delitescens

- Brimley, C. S. Records of Some Reptiles and Batrachians from the Southeastern
United States. Proc. Biol. Soc. Wash. Vol. XXIII, pp. 9-18, March 23, 1910.

3 Cope, E. D. The Batrachia of N.A. Bull. U. S. N. M. No. 34, 1889.

* LeConte, J. Remarks on the American species of the Genera Hyla and Rana. Annals
Lyceum Natural History New York, Vol. I, Part III, New York, 1825, pp. 278-282.

Introduction 5

Thirty years later (1855) he gives 12 forms each of which "inhabits Geor-
gia" or the southern states. The five species which are not yet recorded in
the Okefinokee proper are^

Rana capito Hyla dehtescens

Chorophilus nigrita Scaphiopus sohtarius

Hyla versicolor
In 1832 to 1842 the ten volumes of Dr. Holbrook's epochal "North American
Herpetology'' appeared in two editions. Many of his Georgian specimens or
records came from Dr. Harden of Riceboro, Ga., or from J. Hamilton Coupe r,
of St. Simon's Island. The latter gentleman also knew the country south
of the Altamaha quite well. Dr. Holbrook gives 25 forms from Georgia or
from what might be termed more strictly eastern Georgia. The missing
species from the Okefinokee Swamp in 19 13 are^
Rana pipiens Latreille
Cystignathus nigritus LeConte
Scaphiopus sohtarius Holbrook
Hyla squirella Bosc
Hyla dehtescens LeConte
Bufo americanus LeConte
Salamandra auriculata Holbrook
Salamandra quadrimaculata Holbrook
Salamandra quadridigitata Holbrook
Salamandra fasciata Green
Triton niger Green
Menopoma fusca Holbrook
Siren intermedia LeConte
Siren striata LeConte
Seven years later, 1849, the same author compiled a catalogue of the Batrachia
of Georgia and in addition to the forms given above he enumerates ten more.
They are^

Cystignathus ornatus Holbrook Salamandra biHneata Green
Bufo erythronotus Holb. Salamandra melanosticta L Gibbes

Salamandra guttohneata Holb. Salamandra venenosa Bart.
Salamandra salmonea Storer Salamandra talpoidea Holb.

Salamandra erythronota Green Bufo americanus LeConte
Most of the material which Cope had to examine from Georgia came from
the southeastern or southwestern districts of the state. In the southeast at
Riceboro or in Liberty Co., he had specimens collected by Major J. LeConte,
two forms by Dr. J. L. LeConte, three secured by Prof. S. F. Baird, several
by Dr. W. L. Jones and some not accredited to collectors. In the first group
(of Maj. J. LeConte) occur,^

"LeConte, J. Proc. Acad. Nat. Sci., Phila. Vol. VII, 1854, 1855, pp. 423-431. Phila.
1856. Read Dec. 25, 1855.

^Holbrook, J. E. N. A. Herpetology. Vols. 4, 5. 1842.

' White, Geo. Statistics of the State of Georgia: etc. Savannah. 1849. Appendix p. 15.

8 Cope, E. D. The Batrachia of North America. Bull. U. S. N. M. No. 34. Wash. 1889,
PP- 1-515-

6 Frogs of the Okefinokee Swamp

Chorophilus occidentalis Rana catesbiana

Hyla squirella Rana areolata capito

Hyla versicolor Amblystoma conspersum

Scaphiopus holbrooki Amblystoma opacum

His son, Dr. J. L. LeConte took two forms one of which is absent from the
swamp, to wit, Chorophilus nigritus.

Dr. W. L. Jones collected 14 species of salamanders and three anurans
from the same locahty. The following nine forms are of interest.

Plethodon cinereus cinereus Amblystoma punctatum

Spelerpes guttoHneatus Amblystoma talpoideum

Spelerpes ruber stictoceps Desmognathus fusca

Manculus quadridigitatus Desmognathus nigra
Amblystoma opacum

Whether ''Dr. J. Jones, Georgia" collected or resided at Riceboro we do not
know. He took

Hemidactylium scutatum Stereochilus marginatus

From Riceboro Prof. Baird had three forms, two of which were
Rana virescens virescens Rana catesbiana

Also from Riceboro, Ga. with no collector mentioned Cope hsts six forms,
the four pertinent records being

Hemidactylium scutatum Manculus quadridigitatus

Gyrinophilus porphyriticus Spelerpes bihneatus

Thus we have recorded (until 19 14) to the immediate northeast 21 sala-
manders and 21 frogs, toads and tree toads, while in the swamp we have
recorded, (until 19 14) 5 urodeles and 10 anurans. The forms of Riceboro
missing from the Okefinokee (as recorded until 19 14) are

Ambystoma talpoideum Desmognathus nigra

Ambystoma opacum Notophthalmus v. viridescens

Ambystoma punctatum Pseudobranchus striatus

Ambystoma conspersum Bufo 1. americanus

Hemidactylium scutatum Scaphiopus holbrooki

Plethodon cinereus cinereus Chorophilus ornatus

Plethodon cinereus erythronotus Chorophilus occidentahs

Stereochilus marginatus Chorophilus nigritus

Gyrinophilus porphyriticus Hyla squirella

Manculus quadridigitatus Hyla versicolor

Spelerpes bislineatus Hyla dehtescens

Spelerpes guttolineatus Rana pipiens

Spelerpes ruber stictoceps Rana catesbeiana

Desmognathus fusca Rana areolata capito
Desmognathus fusca auriculata

Introduction 7

To these forms there should be added the three following species from
localities southeast or northwest of the swamp, namely

Rana aesopus Rana areolata areolata

Hyla pickeringii

This hypothetical list from three regions near the swamp probably is far in
excess of what will ever be found in the swamp or its immediate environs.
These 18 salamanders and 13 frogs and toads — 31 in all — emphasized in 1914
the need of further collecting in this region. No doubt some of them hke the
species of Pseudacris (Chorophilus) may be reduced in number and specimens
comparable to both Rana pipiens and R. sphenocephala may be found. We
were on the lookout for members of the Rana areolata group but none of the
three forms appeared. The natives maintain that the bullfrog of the outside
is different from the one of the swamp and the former may represent R.
catesheiana. The two great surprises are the lack of salamanders and the
absence of such forms as Hyla squirella. The author in 19 14 had the convic-
tion that exclusive or extensive collecting of amphibia in this region would
yield several more forms.

In 187 1 and 1876 Paul Fountain visited this region and his amphibian
notes which follow may have enough pertinent historical interest to warrant
their inclusion in this paper .^

"Probably the next hving creature we notice will be the bull-frog, which
attracts us on account of its, to our eyes, huge size. There are at least four
species of frogs in these swamps, including the largest and smallest of American
frogs. The bull-frog is not, however, very abundant, and I have noticed that
this frog seems to prefer clear running streams to swamps. It grows to a very
great size in the Okefinoke swamp, and sitting up on a snag or mass of moss,
it looks like a large stone or boulder. As we approach, however, we notice
the exceedingly brilliant eye fixed steadily upon us, and on attempting to
touch it, it springs away with an alacrity for which we were quite unpre-
pared, considering its heavy, unwieldy build. It has an enormous head, a
thick-set body, and strongly formed hmbs, much thicker in proportion than
those of the common European frog. Its croak is spoken of as "Booming"
and is very loud. It can sometimes be heard appearing to come from every
part of an extensive district, and from a mile or more distant. / am not sure
that the Okefinoke frog is not a distinct variety of the cominon hull-frog. It is
larger than those which I have seen in streams running into the Mississippi
and its tributaries, and in other places, and seems to differ in colour, though
neither of these circumstances is of much importance. Size may depend on
locality and abundance of food, and coloration certainly does. The following
are the measurements of the largest specimen that I could find: Length of
body from nose to base, 8 inches; breadth across, nearly 5 inches; girth, 14
inches; length of hind-limb, 12^ inches; weight, within a fraction of three
pounds. The usual size is about seven inches in length, half as much more

* Fountain, Paul. The Great Deserts and Forests of North America. London, 1901,
pp. 62-64, 67-68.

8 Frogs of the Okefinokee Swamp

for length of hind-limb, and a weight of over two pounds. It is remarkable
that so small a snake as the mocassin can swallow a bull-frog whole. Of
course the snake is greatly distended by such a meal, and is helpless until it
has digested it.

"The bull-frog is said to eat small fish. I cannot say this of my own
knowledge. They feed on slugs, larvae, beetles, flies, spawn, water-insects,
and other frogs, for I saw a bull-frog one day pounce upon a grasshopper-
frog and swallow it in a twinkling, and one that I killed had a dead grass-
hopper-frog in its stomach. I have also found the comminuted elytra of
water-beetles in their stomachs, but no scales or other signs of fish. The
prey is swallowed with great rapidity — in a flash of time.

"The colour of the bull-frog is dusky grey, lighter on the sides and limbs,
and lighter still on the under parts; but the same tinge throughout. There
are some dark spots on the neck and back, and the hind-limbs have leopard-
like markings.

"The grasshopper-frog, referred to above, is very abundant in the swamps.
It is the smallest frog in the States, being only an inch and a half in length,
and weighing less than an ounce. It is greyish-green in colour, with some
brown patches and light stripes on the body, and the legs barred with brown.
There are discs to the toes, like those of a tree-frog, but its habits do not seem
to be arboreal. On the contrary, I have only found it in stagnant swamps
and marshes where there was a thick matting of water-plants. They sit on
the floating leaves in great numbers, and are very agile, and they make a
piping noise, quite unlike the ordinary croaking of a frog. Thousands of
them fall a prey to the storks and other aquatic birds, yet there are myriads
of them in the Okefinoke Swamp.

"In certain parts of the swamp you may see the tadpoles through the
water, swarming on the bottom as thickly as the mosquitoes in the air. I
suppose these to be chiefly the tadpoles of the grasshopper-frogs ; those of the
bull-frogs I could never find.

"The other two frogs found in these swamps are of species which I could
not identify. They are of medium size and dull colour, and there is nothing
remarkable in their habits.

"There are also one or two species of water-lizards — newts apparently.

"Should we dig in the mud near the centre of the swamp, where there is
a sluggish current in the water, it is not improbable that we shall turn up a
hellbender, a species of salamander, which is held in utter abhorrence by our
friends the shingle-cutters. I heard one of these gentlemen refer to me in
the following eulogistic terms : 'Never tell me that that there monkey-crank
ain't in league with the devil. Gawd-a-mighty! I seed him strake a hell-
bender, and never no hurt come to him.'

"The hellbender is as harmless as a frog or toad, or any other causelessly
persecuted reptile. It is an absolutely water-hunting species, which I have
never seen on dry land. It is more abundant in the Mississippi and the great
rivers which run into it than elsewhere, and is scarce in swamps, to which it
seems to resort to bury itself in the mud to hibernate, and perhaps to breed.

Introduction 9

At all events, I have seen the young in the Okefinoke, and they seem to be
hatched from the eggs fully developed, which is different from other sala-
manders, which undergo several transformations before attaining full de-
velopment.

"The hellbender is from fifteen to eighteen inches long, and usually weighs
about two pounds. It has a somewhat eel-like head, a bulky body, and a
flattened tail, adapted to its aquatic life ; but it is not a swift swimmer, and is
easily captured with the aid of a hand-net. The few that I have handled
seemed to be lethargic creatures, and made no resistance to capture, or at-
tempts to bite in which they also differ from most other salamanders. The
colour is grey, with spots and zigzag blotches of darker grey. It burrows in
the mud in search of prey as well as to hibernate, and devours every kind of
slug, worm, fresh-water shrimp, insect, tadpole, or small fish which it can
capture. Its haunts make its habits difficult to study, but I beheve I am
correct in stating that it lies hid in the mud, with only its head uncovered,
for the purpose of surprising the fish as they swim by. I have seen one so
lying with a fish in its jaws. The people of the country have a strong prejudice
against it and never lose an opportunity of destroying it."

The collection of the amphibian material was made at four different
periods. The initial party consisting of zoologists from Cornell University
stayed from May 28 to July 13,1912. It consisted of Professor J. C. Bradley,
the leader of the party, and Professor C. R. Crosby, both of the Department
of Entomology; Dr. A. H. Wright of the Department of Zoology; Principal
W. D. Funkhouser of the Cascadilla Preparatory School; Messrs. M. D.
Leonard, S. C. Bishop, and A. R. Cahn, all of the class of 1913, and Paul
Battle of Bainbridge, Ga., Mr. E. L. Worsham, State Entomologist of
Georgia, and Mr. Chas. S. Spooner were with the party for a week. A
smaller party from the University spent two weeks in the swamp from Dec-
ember 18, 1 9 13, to January i, 19 14. This party consisted of Professors J. G.
Needham and J. C. Bradley, Messrs. Paul Battle and John T. Needham.
Another container was again intrusted to the Lees until August i, 19 14
when it was returned. In addition, the amphibians collected May 6-23,
1 91 2 by Mr. Francis Harper were given to our museum and were available
for this study.

SUBSEQUENT EXPLORATIONS (19 14 TO 1922)
In 192 1 we proposed to enter Okefinokee to make a thorough study of its
vertebrate life, particularly the life histories of the Salientia. With the
support of the Heckscher Foundation for the Advancement of Research,
Mrs. Wright and I planned to leave in April and spend the spring and summer
in the swamp. Circumstances necessitated her presence in Ithaca and Mr.
Francis Harper was secured as my associate. Together from April 21 to
August I we collected in the swamp with Billy's Island as our base. The
Hebard Cypress Company were operating from this island and to them (The
Hebards, John M. Hopkins, A. J. Armstrong, Sam Edwards, Sam Mizzell

lo Frogs of the Okefinokee Swamp

and Hamp Mizzell, The Quarterman brothers, and other officials) and its
numerous employees we extend our heartfelt thanks for numerous kind-
nesses, courtesies, and assistance in our work.

In 1922 we motored to the swamp from Ithaca and arrived June 11, This
year we chose Chesser Island on the eastern edge of the swamp to study the
surrounding region and less disturbed parts of the swamp. On August 2 to 7
we visited Billy's Island. We sought also to get more illustrations of the
breeding stages of life histories we worked out in 192 1 and fill in the gaps in
life histories of species not known.

The party of 1922 consisted of Mrs. Anna Allen Wright, Messrs. F. Harper
and Miles D. Pirnie and myself. With a Ford we were able to operate from
Waycross to Jacksonville or from the island to mainland.

In 192 1 and 1922 we added to our amphibian species of previous years
these species:

Siren lacertina Pseudacris nigrita

Pseudobranchus striatus Hyla squirella

Ambystoma angulatum Rana aesopus

Triturus viridescens dorsalis Rana heckscheri

Scaphiopus holbrookii Rana virgatipes

The frogs in this list are forms solely outside the swamp or mainly so.

THE OKEFINOKEE REGION

The region of this study is roughly the Okefinokee swamp.

The Okefinokee region falls largely in the Sabalian and Lower Austral life
zones of Rehn and Hebbard (1916), or physiographic areas called Lower
Coastal and Upper Coastal by the same authors. It is in the Coastal plain
of the southeastern United States and very decidedly of the pine barren
region variously placed in Flat Pine Barrens, East Florida Flatwoods section
(Florida portion of region). In some interpretations it might fall into the
Gulf Strip of the Lower Austral or in some discussions be thought of as a
part of the northern portion of the range of some "Floridan forms."

The swamp covers parts of Charlton, Ware, CHnch, Pierce, Camden, and
Brantley counties and extends into Florida if Bay Swamp be conceived as a
part of its area. We have done desultory collecting in Nassau and Baker
counties in Florida. In a previous discussion we have given its area as 660
square miles," 26 miles in greatest width and 39 miles in greatest depth.
Possibly these dimensions are somewhat larger than they are now known to
be. The chief effluent of the swamp is the Suwannee river. The St. Mary's
also drains some of the eastern part of the swamp. The tributaries or branches
on the east side of the swamp are mainly short and most of the larger creeks
are tributary to the St. Mary's river, while on the west side and north
many creeks of considerable length flow into the swamp and through it
eventually into the Suwannee. This swamp then makes of east Florida an
island, if one wishes so to term it.

Introduction 1 1

PLANT HABITATS

In 1926 we prepared a paper on The Plant Habitats of Okefinokee Swamp
(MSS) and therein expanded the habitats of the swamp. We will content
ourselves in this discussion with a short general discussion from that paper.

The swamp's geographic subdivisions as given by residents and by the
maps fall into four major groups: cypress bays (such as Billy Island Bay,
Floyd Island Bay, Double-0-Bay, Jackson Bay); prairies (such as Floyd's
Island Prairie, Grand Prairie, Chase Prairie, Honey Island Prairie); islands
("piney islands" such as Billy, Honey, Floyd's and Chesser Islands and
"hammocks" such as Craven, Mixon, Hickory, Cedar); and water-courses,
(a) wooded (such as Minne Lake, Minne Lake Run, Billy Lake, River Nar-
rows, Suwannee River), (b) artificial (such as Suwannee Canal), (c) open
channels through the prairies connecting lakes or water courses. These are
usually made by the residents and often follow the alligator trails. In March
1915 (Phila. Acad. Sci. 1915, pp. 144, 145), Messrs. Bishop and Wright used
these four divisions in classifying the snakes according to habitat.

In 192 1 we used thirteen or fourteen groups for characterizing distribution
of the animal forms within the swamp, namely pine barrens, island edges,
hammocks, grassy fields, cultivated fields, around and in buildings, sand
scrub (Floyd's) cypress bays, cypress ponds, cypress heads, prairies, sphagnum
bogs, and watercourses.

In 1922 being located in Chesser Island near the swamps edge and roaming
on the mainland as we did the list reached 23 or 24 separate habitats or
headings on our plant sheets. Categories such as these were used in gathering
our notes together: St. Mary's river; sand bluff, St. Mary's river; sand scrub;
sand ridges or hills; dry pine barrens; Camp Corneha; cleared and cultivated
fields; around buildings, roadsides and railroads; village streets; Spanish
Creek moist woods; hammocks; moist pine barrens; edge of islands; cypress
ponds; river swamps, branch swamps; bays; strands; edge of wooded lakes;
borders of prairie lakes.

The major natural plant ecological communities or associations of the
swamp proper might roughly be divided into six groups :

1 . "Prairies" (including center of cypress pond, also open lakes).

2 . "Bays" (including edge of cypress ponds, "heads in prairies,"

borders of wooded water courses such as Minne Lake).

3 . Water courses (These merge with "prairies," open-centered cypress
pond).

4. Hammocks (including cultivated fields, buildings, most "old fields").

5 . Barrens (including pine barrens, moist and dry, sand scrub, sand
hills or ridges, etc.)

6. "Strand" (including sphagnum bogs, fern bogs, fern prairie, sphag-

num circle around "heads" and around cypress bases, etc.)

The "prairies" and cypress "bays" are peculiarly distinctive of the swamp.
On the mainland, the open-centered cypress pond suggests these two ele-
ments. The water courses within the swamp can hardly be thought of as

12 Frogs of the Okefinokee Swamp

distinctive botanically as the "prairies," "bays," hammocks and barrens.
The islands of the swamp are hke a piece of the cut-off mainland with its
hammocks and barrens. Witness the mainland extending into the swamp, as.
The Pocket, followed closely by Jones Island, Gallberry Island, and Billy
Island. The "strand" or sphagneous formation in different forms extends
from the pine barrens to the "prairies."

If we proceed from the "prairies" on the eastern edge of the swamp over
Chesser Island to St. Mary's river we have (without consulting notes) these
impressions on breeding habitats based on our 1922 experiences.

Open prairies:

Acris gryllus Hyla cinerea

Rana gryHo

Outer edge of cypress "bay;" open cypress pond in "bay"; edge of lakes
in "prairies":

Acris gryllus Hyla cinerea

Rana grylio

"Strand" of "prairies"

Rana virgatipes

Cypress "bay" itself:

Hyla (in next category)
Three species of "prairies,"
Rana clamitans at island's edge.

Cypress pond on Chesser Island :

Acris gryUus Bufo quercicus

Hyla cinerea Bufo terrestris

Hyla femoralis Rana aesopus

Hyla gratiosa Rana clamitans

Pseudacris ocularis Rana sphenocephala
Gastrophryne caroUnensis

"Dreen" (Cypress "bay" between island and mainland)

Acris gryllus Rana grylio

Hyla cinerea Rana clamitans

Rana virgatipes

Permanent open ponds in pinelands on mainland. (Anna's Pond, Petty
Pond, etc.)

Rana sphenocephala Hyla squirella

Rana aesopus Hyla gratiosa

Hyla femoralis Bufo terrestris

Gastrophryne caroUnensis Bufo quercicus

Ponds (with shrubs or cypress)

Pseudacris ocularis Hyla gratiosa

Gastrophryne caroUnensis Rana aesopus

Hyla femoralis Rana sphenocephala

Imirodudion 13

Roadside low flat temporary pools, no shade :

Bufo quercicus Hyla gratiosa

Bufo terrestris Hyla squirella

Gastrophryne carolinensis Scaphiopus holbrookii

Camp Pinckney pools, high oak banks:

Bufo terrestris Rana sphenocephala

Scaphiopus holbrookii Hyla cinerea

Rana clamitans Hyla versicolor

River swamp (St. Mary's River) pools:

Rana clamitans Rana heckscheri

Et al.

A CATALOGUE OF THE SPECIES OF SALIENTIA OF THE

OKEFINOKEE REGION (1922)

(Plates I, II, III)

The Salientia of this region include species, representing seven genera.
These are Scaphiopus, Bufo, Acris, Pseudacris, Hyla, Rana, and Gastrophryne.

. I. Scaphiopus holhrookii (Harlsin) . The spade-foot.

This subterranean species was found only in the region east of the
swamp. We recorded it from Braganza, Georgia to HiUiard, Florida.
Usually our records are from tadpoles and not from the adults.

2 . Bufo quercicus Holbrook. The oak toad.

This is an abundant breeder in shallow pools during warmer rains. It
may hide during the day or travel abroad in the heath or grassy cover
of the pinelands,

3 . Bufo terrestris Bonnaterre. The southern toad.

This terrestrial species is abundant, being found in almost every land
habitat where cover exists, particularly in pine barrens and hammocks.
At breeding season it seeks shallow water for breeding.

4. Acris gryllus (LeConte) . The cricket-frog.

These little jumpers are in myriads on the prairies and in cypress ponds,
and after rains may be common on the moist islands.

5. Pseudacris nigrita (LeConte). The black chorus frog.

In early spring this species breeds in shallow water not within the
swamp. After breeding it is found on pinelands or high sandy ridges
(Trail Ridge, etc.).

6 . Pseudacris ocularis (Holbrook) . The Httle chorus frog.

This midget of frogs of the U. S. A. breeds in the cypress ponds and
thereafter is usually in the wet pine barrens of the islands' or swamp's
edges, or around the edges of ponds.

14 Frogs of the Okefinokee Swamp

7 . Hyla cinerea cinerea (Schneider). The green tree frog.

The "bellfrog" inhabits waterhly prairies, cypress ponds, swampy
thickets, and proves one of the commonest anurans of such a habitat.

8 . Hyla femoralis Latreille. The pine woods tree frog.

This tree frog is a common inhabitant of the high long-leaved pines of
pinelands and other trees of hammocks except at breeding times when
it frequents cypress ponds and temporary depressions and ditches.

9 . Hyla gratiosa LeConte. The Florida tree frog.

This, the largest treefrog of the region, is scarce within the swamp but
common enough in trees on the outside of the swamp. It breeds in
cypress ponds and other ponds.

10. Hyla squirella Latreille. The Southern tree frog.

This species was not found within the swamp. It normally lives about
domiciles along water courses, and in other habitats in trees, but breeds
in ponds, temporary pools, roadside ditches, etc.

11 . Hyla versicolor versicolor (LeConte). The tree toad.

We found this common species the rarest Hyla in this region. We took
it only along the St. Mary's River. Here it bred in pools under oak
trees.

12. Rana aesopus (Cope). The gopher frog.

This inhabitant of gopher turtle holes along the Trail Ridge repairs
to cypress and other ponds for breeding.

13 . Rana clamitans Latreille. The green frog.

This species is considerably smaller than in the north, less common
than several Ranas, and frequents the edges of cypress ponds, cypress
bays, border of streams, etc.

14. Rana grylio Stejneger, The southern bullfrog.

This species is common on the prairies, along the edges of open water
lakes, or in openings in the cypress bays.

1 5 . Rana heckscheri Wright. Heckscher's frog.

This species described from this region is an inhabitant of river swamps,
swampy tangles outside Okefinokee swamp proper. In these places it
breeds.

1 6 . Rana sphenocephala (Cope) . The southern meadow frog.

This is the common spotted frog of the region, breeding in cypress
ponds, cypress bays, and ponds of other description as well as streams,
runs, dreens, etc. Later it travels more on the land in moist cover.

1 7 . Rana virgatipes Cope. The carpenter frog.

This species was not common in the center of the swamp. In the
cypress edge of Black Jack Island and in the cypress bay edges of the
east edge of the swamp, and in the prairies in the nearby region we
recorded it. It breeds in the cypress bays.

Plate I

Lateral aspects. Bufo, Gastro'phryne, Pseudacris, Scaphiopus

1. Bufo americanxis, male, Ithaca, N. Y., May 4, 1929. From W. J. Koster.

2. Bufo fowleri, female, Mt. Vernon, Va., Mr. and Mrs. M. C. Brady and author

June 3, 1928.

3. Bufo lerrestris, male, Gainesville, Fla., May 4. 1929. From O. C Van Hyning.

4. Scaphiopus holbrookii, male, Mobile, Ala., Apr. 18, 1929. From H. P. Loding.

5. Bufo quercicus, male, Gainesville, Fla., May 7, 1929. From O. C. Van Hyning.

6. Gaslrophryne carolinensis, female. New Orleans, La., May 25, 1928. From

Percy Viosca, Jr.

7. Pseudacris ocularis, Billy Id., May 20, 192 1.

8. Pseudacris ornata, male, Helotes, Texas, Feb. 22, 1925.

9. Pseudacris nigrita, Chesser's Id.. July 16, 1922.

w^

V^Sijiif.

Plate II
Lateral aspects. Hyla and Acris
Acris gryllus, female, New Orleans, La., May 19, 1928. From Percy Viosca, Jr.
Hyla crucifer, female, Ringwood, Ithaca, N. Y., Apr. 7, 1929.
Hyla andersonii, male. Cape Ma_v, X. J., June 1928. From Dr. and Mrs. A. B.

Klots.
Hyla graliosa, male, New Orleans, La.. Apr. 30, 1928. From Percy Viosca, Jr.

5. Hyla cinerea, male, Gainesville, Fla., May 20, 1929. From O. C. Van Hyning.

6. Hyla evittata, male, Mt. Vernon, Va., Mr. and Mrs. M. C. Brady and author,

June 3, 1928.

Hyla squirella, male. New Orleans, La., May 19, 1928. From Percy Viosca, Jr.

Hyla femoralis, female, 8t. Petersburg, Fla., Aug. 7, 1928. From X. P. Fry and

Son.

9. Hyla versicolor, male, Pt. Henry, X. Y., June 17, 1929. From Carl Van Deman.

[o. Hyla avivoca, male, Xew Orleans, La., May 19, 1928. From Percy Viosca, Jr.

<#*■

••-^

lO

Plate III
Lateral aspects. Rana males

1. Eana septentrionalis, Onekio, X. Y. July 14, 1923. x i.o

2. Rana damitans, Billy Id., Ga. May 23, 192 1. x 0.8.

3. Rana virgatipes, Chesser Id., Ga. July 22, 1922. x i.i.

4. Rana heckscheri, half-grown adult, Hilliard, Fla. August 17, 1922. x 0.75.

5. Rana aesopus, Chesser Id., Ga. 1922. x 0.65.

6. Rana pipiens, Vermont. Oct., 1928.

7. Rana sphenocephala, Billy Id., Ga. May 18, 192 1. x 0.5.

8. Rnnn gri/lio, Billy Id., Ga. May 9, 1921. x 0.8.

General Discussion

15

18. Gastrophryne carolinensis (Holbrook). The narrow-mouthed toad.
This subterranean species becomes abundantly evident when suitable
rains occur. Then its sheep-Uke bleat is heard frequently in the pine
barrens, edges of hammocks, etc.

HYPOTHETICAL SPECIES TO BE EXPECTED OR
SOUGHT IN THE OKEFINOKEE REGION

In view of the foregoing discussions there is more or less (sometimes
remote) likelihood of the following species being recorded near or in the
Okefinokee swamp.

Bufo fowleri Hyla andersonii

Pseudacris occidentalis Hyla avivoca

Pseudacris ornata Rana catesbeiana

Rana pipiens (see R. sphenocephala)

In this paper the life history of Hyla andersonii is considered and a few
observations passed on Pseudacris occidentalis and Pseudacris ornata. The
life histories of the common forms Bufo fowleri, Rana catesbeiana and Rana
pipiens are omitted. For comparison with Rana virgatipes, the life history
of the northern mink frog (Rana septenirionalis) is incorporated.

GENERAL DISCUSSION OF THE OKEFINOKEE
SALIENTIA

RANGE

It would be natural not to expect truly tropical Floridan species such as
Scaphiopus holbrookii albus or Eleutherodactylus ricordii (West Indies — Gaines-
ville, Florida) in the Okefinokee swamp, yet on the Keys and in this southern
tropical region such Sabalian forms as Hyla squirella and Hyla cinerea and
Bufo quercicus occur.

The Sabalian frogs comprise the bulk of the swamp's amphibian inhabi-
tants. About eleven species occur mainly in the Sabahan region. They are
all, with two exceptions, in the Swamp (I.O.) or outside (0.0.) The two
exceptions may later be found when better understood.

0.0.

Hyla squirella

Va. -

-Tex.

I.O.

Hyla femoralis

N.C.-

-Fla.-

-Tex.

I.O.

Bufo quercicus

N.C.-

-Fla.-

-Ala.

Pseudacris occidentalis

s.c-

-Tex.

Pseudacris ornatus

s.c-

-Tex.

0.0.

Pseudacris nigrita

s.c-

-Fla.-

-Miss.

I.O.

Hyla gratiosa

s.c-

-Fla.-

-La.

I.O.

Pseudacris ocularis

s.c-

-Fla.

0.0.

Rana aesopus

s.c-

-Fla.

0.0.

Rana heckscheri

s.c-

-Fla.

I.O.

Rana grylio

Ga.-

-Fla.-

-La.

1 6 Frogs of the Okefinokee Swamp

Four of these are solely outside and Hyla gratiosa mainly so. Rana
aesopus is absent within the swamp because of its subterranean habits.

There are about nine species of frogs which might be considered Austral,
Three of them are absent such as Bufo fowleri, Pseudacris feriarum and
Hyla andersonii. The latter ought to be present since its associated species
Rana virgatipes occurs in the swamp. The six Austral species are

Bufo fowleri

N.E. — Ga.-

-Tex.

0.0.

Scaphiopus h. holbrookii

Mass. —Fla.

I.O.

Acris gryllus

N.Y. — Fla.-

-Tex.

I.O.

Hyla cinerea

N.J. — Fla.-

-Tex.

I.O.

Rana virgatipes

N.J. — Ga.

Hyla andersonni

N.J. —S.C.

I.O.

Gastrophryne carolinensis

Va. — Fla.-

-La.

I.O.

Bufo terrestris

S.C. — Ga.-

-La.

The widespread Hyla versicolor was recorded on St. Mary's River at
Folkston, Ga. and Rana catesbeiana has been at times reported to be near
the swamp though we did not take it.

COLORATION OF SPIRIT SPECIMENS (191 2)
AND MEASUREMENTS OF 191 2 MATERIAL

These are parts of the 19 13 manuscript. They were painstakingly made
and are herein included as contributory to an understanding of the species.
They can be interpreted in part as more pertinent to the 1912-1914 period
in amphibia of the U. S. A. but in part they are apropos to the present dis-
cussion and paper, because partially they pertain to species little understood
and partially because they were conceived from a different angle than the
coloration from life or the scheme of measurements (relative) especially em-
ployed in this paper.

MEASUREMENTS

We have made relative measurements of the Okefinokee frogs to accord
with a system we are employing for all the frogs of the United States. We
choose from our and U. S. National Museum collections (supplemented from
the collections of other institutions) representatives of each species and
subspecies which measure in body length 20 mm., 28 mm., 36 mm., 44 mm.,
56 mm., 66 mm., 68 mm., 82 mm., 95 mm., 108 mm., 125 mm., 136 mm.,
and 150 mm. In this way if one of each of the Ranas of the swamp are meas-
ured at the same body length we have all our actual measurements in part
also relative if they be put in a table. For example at 56 mm., such a table
as the following might be constructed. (Table i)

General Discussion

Table i

Number

I

2

3

4

5

6

7

8

Ratm

virg.

sept.

clam.

heck.

gryl.

spheno.

aeso.

cates.

Sex

d^

&

c^

im.

im.

&

im.

im.

Length (sn.-vent)

56

56

56

56

56

56

M

56

Head (tympanum)

21

24

26

2i

21-5

23-5

i

22

Head (angle mouth)

15

19

19-5

20

20

21

s

20

Width of head

18

23

24

24

21

21

s

23

Snout

8

9

10

10

10

10.5

i

9

Eye

6.5

7

8

7

6.5

7

n

7

Interorbital space

2

3

35

3

3-5

4.0

g

4

Upper eyelid width

4-5

6

5-5

6

5

6

4-,

Tympanum

7

10

9

5

5

6

i

5

Intertympanic width

12.5

II

12

iS-5

12

14-5

n

15

Internasal space

4-5

4

6

S-5

3-5

5-5

6

Forelimb

2i

30

31

29

27

30

t

31

ist finger

7

7

9

9

9

9-5

h

10

2nd finger

7

9

8

7-5

9

7

i

10

3rd finger

9

10.5

12

10

13

II

s

12

4th finger

6

Q

7

7

10

7

10

Hind limb

75

88

86

84

71

104

s

82

Tibia

23

29

31

29

36

e

28

Foot

25

32

33

30

25

33

r

28

ist toe

7

7

8

8

9

6

i

9

2d toe

13

13

12

12

13

13

e

13

3d toe

18

20

18

17

18

18

s

18

4th toe

24

28

30

27

25

30

27

5th toe

16

19

20.5

18

23-5

20

20

In this paper we will not attempt to include all the measurements for the
species of the Okefinokee region. These are reserved for a growth paper in
process of organization and therein will be incorporated.

This chart means that if one finds a specimen 6-10 mm. in body length
it might be a transforming or transformed frog of nine species; or at 11-20
mm. it might be one of 16 frogs (one of 15 transforming species or young of
14 species, or a male of 6 species or a female of 5 species) ; at 68 mm. there are
ten species to consider, a male or a female of Bufo terrestris, Scaphiopus
holbrookii, Hyla gratiosa, Rana sphenocephala, Rana septentrionalis, or Rana
damitans, a male of Rmia aesopus, or immatures of Rana heckscheri, R.
grylio, or Rana catesheiana; at 95 mm. four species only are to be considered,
a female Rana aesopus or either a male or a female of R. heckscheri, R. grylio
and Rana catesheiana; at 150 mm. only two species enter, a male or female
each of R. grylio and Rana catesheiana.

This chart might suggest the desirability of a key at each size for true
comparisons. At 6-10 mm. it might thus prove a more desirable key because
each of the nine species are at transformation and are comparable. In the
same way at 95 mm. we have mature males and females of the green frog,
gopher frog, Heckscher's frog, southern bullfrog, and bullfrog. Even here
however we have the green frog and gopher frog nearing the extreme of their
size and females of the other three just about beginning breeding. Possibly

Frogs of the Okefinokee Swamp

a s
s s
s s

cl « -^ .S '^ '^ ci ■-

^ Jill § -S^ S . I S|-|^-'i.^^|.t| I §1,1

B pqpQccffi WKffi WKP^p:^^ p^ p:? pc; p^ £

General Discussion

19

old age maximum frogs might not be comparable. In the same way if one
glance at sizes from 11-20 mm. to 82 mm. one will find at any one size differ-
ences in maturity. Nevertheless keys at each size might prove useful and
might in spite of their limitations prove suggestive of better characters than
those we have chosen more at random in the past. In this connection, how-
ever, the keys of the past have not been so bad because they have generally
chosen breeding adults. Possibly they will prove as good as some suggested
by this identification index. (Table 2)

We have sought to measure the Salientia (frogs, toads, tree-frogs, etc.)
of the United States and Canada at the 13 sizes given under the foregoing
topic (see Measurements). Our intention was to present in this paper a
series of keys at these various measurements. This will be delayed until
the missing sizes can be found. Any student of the frogs of the Southeastern
United States may find the following tables at 28 mm. (Tables 3, 4), 44 mm.
(Table 6), 66 mm. (Table 7), 82 mm., 95 mm. (Table 8), 113 mm., 125 mm.
(Table 5), and 150 mm. (Table 9) of use and they are included though our
studies of these measurements and growth are not completed.

Table 3

£ M)

"B

'c

0

0

0

C

M

'E

'C

-S

cr

cr c3

Length 28 mm.

p.;

ai

a^

d

d

<

<

a

a

a

«

m K

Sex

c^

9

&

&

9

cf

9

c^

9

im

cf

9 im

H. tympanum

10

II

10

8.5

9.0

H. ang. mouth

6

5.6

7-5

9

8-5

7

6

Width head

8

8

10

9-5

12

10.5

9 5

Snout

4

3-4

5

5

4 5

4

4

Eye

2.2

32

28

4 5

4.0

3 5

30

Interorb. sp.

2 .2

3-2

1.6

3

3 5

3-5

30

Upper eyelid

1.6

1.8

1.6

2.5

3 5

2.5

2.5

Tympanum

—

—

2

3

3 5

2

2

Int. tymp. sp.

—

—

8

8

9 5

8.5

8

Int. nasal sp.

2.2

2.8

2.8

3

2.5

2.5

2.5

Forehmb

12

II

15.8

16

15-5

14 5

14

I St finger

2.0

2.2

4

3 5

3 5

3

3

2nd finger

2.6

38

4

3-5

3

3 5

3rd finger

50

56

7

5

6

5

5

4th finger

4

3-8

4

4

3-5

3

4

Hind limb

2S

26.4

45

34

27

26

26

Tibia

10.4

10.4

17 4

12

II

ID

10.5

Foot with tarsus

16

15 4

21.6

18

15

13

15

Foot without tarsus

II

13 6

II

9 5

10

10

I St toe

2.8

3

5

4

3

1-5

15

2nd toe

4.6

50

7

6

5

3-5

3 5

3rd toe

8

7.8

II

8

6

5

6

4th toe

II

II

13 6

10

8

7-5

9

5th toe

8

7

10.5

8

5-5

4-5

5

Frogs of the Okefinokee Swamp
Table 4

03

(1

P-

tc 0

rr,

1

^

1

.23

0

0

2

1

0^

litani
pus

■5

■3

s

g

2

"S

-C

£? «

lair
eso

O"

cr

OJ

ai

b

c

.3 0,

ai

tc

t4-

ij

>

bC

> ^

0 03

Length 28 mm.

w

W

X

K

ffi"

W

ffi

P^

pi pi

P^ tf

Sex

d"

9

<T

9

im.

im.

im.

ti

t ti

t t

H. tympanum

10.5

10

10

10

10

10

II

12.5

II. 5 *

13 13

H. ang. mouth

9

8.5

8

9

8

9

10

10.5

10

10 11

Width head

10

10.5

10

10

ID

12

115

10

10

II 12

Snout

5

5

4-5

5

5

5

5

5

6

5 25

5 6

Eye

4

3

3

3

3

4

4

4

4

4 5

Interorb. sp.

3

3

3

3

5

3-5

5

3

2.5

2

3 3-5

Upper eyelid

3

3

2

2

5

3

2

5

3

3

2

2.5 3

Tympanum

2

2

2

2

5

2.5

3

2

3-5

2-5

2.5 3

Int. tymp. sp.

8.5

8

9

8

5

8.5

9

9-5

8

8

8.5 10

Int. nasal, sp.

2

3

2

2

5

2

3

2-5

3-5

3

3 4-5

Forelimb

13

135

13

15

155

20

16

15

16

15 15

1st finger

4

3-5

4

4

3-5

3-

5

4

4

4-5

4-5 5-5

2nd finger

4

4

4-5

5

4 5

5-

0

4 5

3-5

4

4-5 5-5

3rd finger

5-5

6.5

6.5

6

5-5

8

6.5

5-5

6

6.5 7

4th finger

5

4-5

5

5

5

50

6

50

4

5

4-5 5

Hind hmb

39

38

42

41

47

44

44

41

44

44 41

Tibia

15-5

—

14 5

15

5

17

16

17

15

13-5

15 —

Foot with tarsus

18

20

20

1 8 20

Foot without tarsus

II

II-5

115

10

115

14

12.5

13

13 14

1st toe

2

4

3

3

5

3-5

3

3

2.5

4

4 3-5

2nd toe

3-5

5

5

4

6

4-

5

6

5-5

6.5

8 6

3rd toe

7-5

8

6

8.5

8

9

7-5

9

9 7-5

4th toe

9

9 5

10

10

ID

II

II

II

12

13 14

5th toe

7

8

6

8

8

10

7

8

8.5 9

*U. S. N. M. & C.

U. collections have 2(

),

30 mm. s

pe

cimen

s not

it .28 mm

Table 5

^

^

bis

M)

Z Z

jz

btj

bC

0

«

pi

rt

pi

P^

^ pi

pi

rt

pi

pi

P^

Sex

&

9

&

9

cf 9

9

&

9

cf

9

Length

113

113 113

113

113 113

125

125

125

125

125

H. tympanum

44

44

39

40.5

43

46.5

42

45-5

43

H. angle mouth

40

39

38

36.5

41

40

39-5

41 -5

41

Width head

48

43-5

38.5

41

47-5

46

42

49

45

Snout

17

16

15

17

19

16

17

18.5

17-5

Eye

12.5

10.5

12

135

13

13

12

12.5

Interorb, sp.

7

6

4-5

■J

8

5

4

8

6

Upper eyelid

9 5

8

7-5

7

9-5

9

8.5

8.5

8

Tympanum

17

19

12.5

12

II 5

20

12

15

II 5

Inter-tymp. sp.

9

175

20

23 -5

26

20

25

24

27

Inter-nasal sp.

9 5

7-5

7

9 5

9 5

6

8.5

8.5

9

Forelimb

45

52

57

61

56

57

62

45

67

1st finger

17

21

18

21.5

19

20

23 -5

21

2i.5

2nd finger

16

24

22.5

22>

18

23

245

21

235

3rd finger

21

28

28,5

27

24

28

30

29

31

4th finger

17

23

23

24- 5

20

23

245

22

255

Hind limb

160

153

145

169

160

168

173

168

180

Tibia

56

54-5

52

56

58

58

55

61

63

Foot with tarsus

78

80

77

82

81

84

86

875

97

Foot without tarsus 52

58

52

58

55

60

60

59

63

I St toe

18

235

23

22

20

26

25

22

23

2nd toe

27

?>i

33

31

28

36

34

32.5

34

3rd toe

38

46

44

44

42

52

47

46.5

49

4th toe

52

58

52

58

54

61

60

59

63

5th toe

38

46

44

41-5

40

51

48.5

46.5

49

General Discussion

'BU'BI9qS9:>'B0 -jj ^

\y-) lO lO I/-) u-) i-O lO

O

•lOlO LOly-jjj-) LOLO ly-)U->

6^ ro r^
ro <N M M oi M

sndosaB •'jj g
sireuou^u8:>das -y; gvo-+i/^t^ioc^ro4MTf-r^coooMoor}-'^<Ni/^ON'i-t^ioco

sadi^^SaiA y;

o

.— Mi_M H-lCS t-,<^(N i_,(Nl-(

ly-J 1/-) U-> LO U-> U->

JOTO01SJ9A -H rv, to .o .o lO lo

tOLOlOVOtOU", VOIO to

• -ti (D^Mroso'tOTl-ro<Ni-iTl-rO\Or^O^OOro'~^ OOioI>.(N\Ow

to to to

On t^ to CO ro <^ c^

nuosjapu-B jj r^

to to to to to to to

TI5[OOjqjOtf 'g r; f^voo^oo'vOO tOTfvD -^^ totooo" •^toiO'+MD <N

O <N t^

to toto totoioto to toto

SIJ^S8JJ9^ 'Q S toTj-c>r^O 't'i-rO'+fO'"' totot^totovo cs rfrotoo^'^oo

?: 33—-! rvX^oj-^ '^3

O O

o o

Frogs of the Okefinokee Swamp

to u-> ir>

OITitjS "Vr pLoo)TfC-c>Oro»or~-rD^(NO«CNOC^t^| roO<NCC-1-0-t
IJ8l[0SJ|09q -jj g

lO to lOtotOLO to toto

sndoss'B yx c'Ovocor^coiotoioot^ovoo^i-''N i roccoi^^ooo

"■.„(NCS(NM <N Mm«»hmCO|t1-Mwm(N<Nm

SU'B^IUI'BIO -U o

lOlO loto lOlOlO lOto

SU'BIIUI'BTO "Vr Pv-v'^^i^OO^iJ^OfO^OO-'Ni-iioi-iioioOOrOMt^-t-rorc
*■ 1 O- U(»(N(Ni-i wi-i <NMi-<i-ii-(C>rorfro>-i"<NfOtN

lO VO to '^ lO to to

•' •*+ <l^-^(NMiN « (NMMMMaro-i-rO"i-CNroO)

to toto tototo to to to

siiBuouiuaidas yi i^^+oioc'corotooot^^oowr^r^'oo
sedp'BSjiA -jj ^

'Bi'Bqdaoouaqds -y; q^

a ^

•i:^ BTBUdaOOUaudS •"Vr rs_r^<^to<NC«0 ^\0 r--r^O to<N0C m t^oo Tt l OCOO POO t^<N

ffl '^ T a-0(N<NOM 1-1 CO" " «t|fO wCNrOIN

<; M

7^

to to to to l'^

'BSOII'BjS 'TJ IV,f<i-i'*<^'^COtovOO^OiJ^i-i(NtOM<Nro| ONOOfNCfOt^

•■' nCJrgc-jCNi-i hi fO"Wl-il-lO"fO|f^ Mr-MM

to to lO

UNOOJqiOU -Q pj,<N00'i-0c0C0O^0MiowvOl^i-iO<N0<^'N^cC<N<^M

toto tototOLO to tototo toiOl^l^

lIJfOOjqTOlI "Q lv.-^OioOr^cOiOtorovOOr^:^'-'OTrwrO'+'+cC!^0~«
•■' "^^Umcscnm cn ro 1-1 >Ocsro<N ^iwi-i

to "^

STJ1S9JJ81 a r^4-^Ot^C^O-f-■>0"^l-l»or^Oa'CoaOTl-I--losoalO<ri<N

to to to to to to

to to to to

6 i..

3 O

o

=3 a-T3

^ c^MW^cgw^tS^H^^fe 2 S ^^WHfi;^^ '^ g ^

CO -^ lO

General Discussion 23

ic 10 i/^ iTi m iT-, m

« CO.. , ^ . _„

sndosaB 'H o eg ^l ^ ^. ^T' "° '^^ ^ "^ ? "" p: IT" ^'

0 fo

0 <N CO ^^ \0 0
Tl- « " IN rO fO

^ ^

00 ir^ -^ Ov Tt- "0
■<^ "H 0) 0) Tj- rO

sub:^™^^ "H '^3 »

m C 0430 ict^rot^t^

10 ITl

llll^ lfellill|||=,'^-1slssg

24

Frog 8 of the Okefinokee Swamp
Table 9

^ J2 J

bC

M

h

0

M

b£

-

r*

u

0

0

pi

d

pi

p4

^

C^

^

pi

pi

pi

pi

Sex

&

9

&

9

&

9

d^

9

d^

9

9

Length

136 1

[36

136

136

150

150

150

150

180

180

184

H. tympanum

50

51

53

44-5

57

48

61

63

H. angle mouth

41

49

51

43-5

53

47-5

57

63

Width head

48

56

47-5

46

56

52

70

73

Snout

18

21

18

18

20.5

21

25

31

Eye

13 5

13

13 5

12

14

14

15

18

Interorb. sp.

5-5

6

6

6

10

6.5

10

9-5

Upper eyelid

9

II

9

9

10.5

10.5

12.5

12

Tympanum

22

18

13

13

22

13

22.5

19

Inter-tymp. sp.

20

28

26.5

255

29

29

32

40

Inter-nasal sp.

7

10

9

7

10.5

9 5

12.5

12

Forelimb

52

66

71

76

70

74

90

80

1st finger

22.5

20

22

22

28

26

35-5

32

2nd finger

24 5

19

22

26

27

26

38

32

3rd finger

32

26

i2

31

34-5

31-5

48

40

4th finger

23 5

20

23

26.5

30.5

27

42

32.5

Hind limb

187

190

165

175

218

173

260

220

Tibia

63

66

57-5

61.5

73

61

92

82

Foot with tarsus

' 93

96

89

98

108

98.5

127

120

Foot without

tarsus

65

—

60.5

66

73

66.5

94

75

ist toe

25

15

22

26

27

27

37

29

2nd toe

39

28

33

40

41 5

38

52

44

3rd toe

50

41

45

53

52

52

72

64

4th toe

65

60

60.5

66

73

66.5

94

75

5th toe

50

42

42.5

53

52

51

70

60

FIRST APPEARANCE AND AUTUMNAL DISAPPEARANCE

These two topics we can discuss in no way clearly or specifically because
we were not in the swamp the last of September, October, November and
December before the Christmas season, nor in the swamp from January to
April 22. It is therefore apparent why we have no reason for a general dis-
cussion of these topics or hibernation. The U. S. National Museum collec-
tions have contributed some records for the above four months.

BREEDING SIZES

On the basis of least breeding sizes we have six groups in the 2 1 species
considered. First come two forms, Pseudacris ocularis and Acris gryllus,
which mature at 11.5-15 mm. for males and 12-16 mm. for females. The
next group of five, namely Bufo quercicus, Gastrophryne carolinensis, Pseu-
dacris nigrita, Hyla squirella and Hyla femoralis, matures for males at 19-

General Discussion 25

24 mm., and for females at 20.5-23 mm. The third group of Umr—Hyla
andersonii, Hyla versicolor, Hyla cinerea, Rana virgatipes, matures for males
at 30-41 mm., for females at 38-41 mm. The fourth group of six species, viz:
Rana septentrionalis, Hyla gratiosa, Rana sphenocephala, Bufo terrestris,
Rana clamitans and Scaphiopus holbrookii matures for males at 48-54 mm.
and for females at 48-56 mm. The fifth group has one form, Rana aesopus,
with first males at 68 mm. and first females at 77 mm. The final and sixth
group comprise Rana heckscheri, Rana grylio and Rana catesbeiana with
males maturing at 82-85 mm. and females, at 85-102 mm. The above data
and more are summarized in the following table: (Table 10)

Table 10

M

ALES

Females

Pseudacris ocularis

II.

5- 15.5mm.

12

- 17.5 mm,

Acris gryllus

15

- 29

16

- 33

Bufo quercicus

19

- 30

20.

5- 32

Gastrophryne carolinensis

20

- 30

22

- 36

Pseudacris nigrita

21

- 28

22

- 30

Hyla squirella

23

- 36

23

- 37

Hyla femoralis

24

- 37

23

- 40

Hyla andersonii

30

- 41

38

- 47

Hyla versicolor

32

- 51

33

- 60

Hyla cinerea

37

- 59

41

- 63

Rana virgatipes

41

- 63

41

- 66

Rana septentrionalis

48

- 71

48

- 76

Hyla gratiosa

49

- 68

50

- 68

Rana sphenocephala

49

- 78

53

- 82

Bufo terrestris

51

- 74-5

56

- 82

Rana clamitans

52

- 72

58

~ 75

Scaphiopus holbrookii

54

- 72

50

- 71

Rana aesopus

68

-lOI

77

-108

Rana heckscheri

82

-113

102

-125

Rana grylio

82

-152

85

-161

Rana catesbeiana

85

-180

89

-200

The relationship of egg diameter to smallest breeding size in each species
is given in the following table (Table 1 1). With the smallest and largest egg
vitelline diameters we divided the least breeding size of each species to see how
many times bigger the body length was than the real egg diameter of each spe-
cies. For example the smallest breeding oak toad in body length is only 13.6
times longer than the egg diameter of that species, wherein the smallest breed-
ing bullfrog would be 50 times longer in body length than its own egg.

The five smallest species have relatively the largest egg vitelli, 13. 6-1 5
times in body length. The bulk of the species (15 in number) have vitelU in
body length from 17.5-30 times. Rana grylio has it 41 times and Rana
catesbeiana has it 50 times.

^3 •

6-15.

8

14-

4-19.

0

15

-16

17-

5-28.

2

i8.

3-20

IQ.

1-28.

7

21 .

4-25

22.

7-27.

3

23-

-28.

7

23-

1-46.

2

24-

5-3 7-

7

25

-36

26.

6-29

27-

2-49

28.

3-37-

7

29

-37-

I

3°

-36.

9

30

-42

41

-58.

5

50

-70.

8

26 Fro^s 0/ </?e Okefinokee Swamp

Table i i

Bujo quercicus
Pseudacris ocularis
Acris gryllus
Pseudacris nigrita
Gastrophryne carolinensis
Hyla Jemoralis
Hyla andersonii
Rana virgatipes
Hyla squirella
Hyla cinerea
Rana sphenocephala
Scaphiopus holhrookii
Hyla versicolor
Hyla gratiosa
Rana aesopus
Rana clamitans
Rana septentrionalis
Bujo terrestris
Rayia grylio
Rana catesheiana

VOICE

The species of Bufo, Hyla, Acris, Pseudacris, Scaphiopus have single
vocal vesicles beneath the chin. The frogs go from single mental vocal
inflation of Rana clamitans, R. heckscheri, R. catesheiana, R. grylio, R. septen-
trionalis (slightly lateral as well), to round paired sacs of Rana virgatipes,
elongate lateral sacs of Rana sphenocephala, to the side of body inflated as in
Rana aesopus.

We will not attempt a general discussion of voice. It is a hard topic to
characterize and the process of voice production in detail is a subject we plan
to pursue at greater length in subsequent investigations.

MATING

Every species (/?. heckscheri and P. nigrita not studied) in Okefinokee
swamp doubtless mates or breeds more at night than by day, but inasmuch
as heavy rains are most responsible for onset of breeding many of them may
mate vigorously by day. Those we have thus found are : Scaphiopus holhrookii,
Bufo quercicus, Bujo terrestris, Acris gryllus, Hyla femoralis, Hyla gratiosa,
Rana aesopus, Rana clamitans, Rana sphenocephala.

The mating eagerness, or ardor has been much discussed as to the mutila-
tion which males will endure when the clasping impulse is paramount. This
we will not discuss in this paper. It is hard to measure which are the more
tenacious — toads, frogs, treefrogs or narrow-mouthed frogs. All are examples
of intense amplexation.

At this point I wish to observe that contrary to some statements that the
extrusion of eggs usually occurs only after the male has clasped the female
for several days, in the species of the United States this seldom obtains. We

General Discussion 27

have seen mated pairs of at least 32-35 species and of some species many
pairs. In general in nature the eggs are laid soon after amplexation. The
females do not approach breeding grounds or males until ready or nearly
ready. Occasionally or rarely a pair may be mated in toads while en route
to pond, but normally they meet at the breeding grounds. Many times
with laboratory mating or sometimes with pairs brought into capitvity several
days elapse before ovulation and then often the eggs do not develop.

We have also seen little indications of unmated males fertilizing the eggs
or egg masses already laid as has been recorded for European forms. One
wonders how many of those observations were made in the field. Further-
more the unmated males will hang about the egg masses laid because usually
this is where the non-ovulated females will lay. They often hide under the
masses, about them, etc. In such areas we hunt for the frogs, males, females
and mated frogs.

If the supernumerary males have accidentally or designedly fertilized egg
masses it has not been observed by us. Query also has been asked what
stimulus prompts the male to discharge his sperm thus without the female.
If they do it maybe the same that has caused individual females of many
species to ovulate without a male or ovulate if the male is removed. Possibly
with some males as with these females the process has gone so far and the
frenzy or intensity of mating so vigorous no stay is possible.

Fall or late summer copulation has engaged some workers' attention. In
the northern states where the ovulation periods of species are short or in
European species some assert that a male will copulate a second time. Whether
it be a second time or a male which ripened late is not always ascertained. If
one glance at the breeding seasons of these Okefinokee frogs as with desert
species, he will discover some breeding from February or March to October i,
November i, or December. This probably means that there are ripening
individuals ready for each suitable rainy period, not necessarily one short
crestal period as occurs in some northern species and that a male or female
breeds only once that season.

Many of us in the north have been accustomed to thinking when we heard
a common meadow frog croak in August or September or a peeper in Septem-
ber or October, that this is an exuberant salutation before it goes into hiberna-
tion. Possibly in some cases it means nothing. In other cases it may mean
males which have ripened. For example, in September 1929 our automobile
went bad at night before a farm in lower Wabash valley. The instant we
got out we heard an immense chorus of southern meadow frogs, Rana spheno-
cephala. We asked ourselves if it were a play chorus. The next day on our
way to Olney, 111. in some ponds we found several fresh masses of eggs. The
fall croaking then in some instances in the north may mean breeding or
incipient urges to breeding.

COLORATION FROM LIFE

In this discussion we have attempted to give a Ridgway description of
live Okefinokee representatives of males and females. Normally descriptions

28 Frogs of the Okefinokee Swamp

have been from spirit specimens. Furthermore the date of the description is
given. A July description might be quite different from a March or an April
one in color tones. True, we could not attempt to catalogue all the variations
in color and these descriptions are only as representative as the season and
circumstances would permit. In almost every species there were pronounced
differences between the males and females of a species.

STRUCTURAL DIFFERENCES

In all the frogs (Rana) the enlarged thumb is the best criterion of maleness.
None of these Ranas of Okefinokee have two excrescences on the male's thumb
as in some of the Pacific coast species of Rana, and none seem to have the webs
of the hind feet as in some northwestern and northern species. In the southern
meadow frog and gopher frog we have not the distinctive differences, of
tympana found in the green frog, carpenter frog, mink frog, Heckscher's frog,
southern bullfrog or common bullfrog. There may be more yellowish tints in
the throats of those five in the males than in the females.

In the Hylidae of the Okefinokee region the series may be separated by
the different color on chin and throat except possibly at times in the pigmy
Pseudacris ocularis. In the same way it is not always easy to separate a male
narrow-mouthed frog from a female of the same species. In the solitary
spade-foots the throats, excrescences on fingers and coloration of the back
generally assist. In the toads the customary differences more or less obtain.

THE CUSTOMARY AMPLEXATION

(PI. IV, Figs. 1-4; PI. V)

In the two toads (Bufonidae) oak toad and southern toad the embrace is
axillary or at times supra-axillary with the first two fingers folded back. In
the nine species of tree frogs ( Hylidae) herein considered, namely, the little
chorus frog, black swamp cricket frog, cricket-frog, southern tree frog, pine
woods tree frog, Anderson tree frog, green tree frog, common tree toad, and
Florida tree frog, the embrace is axillary or supra-axillary. In the narrow-
mouthed toad (Brevicipitidae) it is axillary. In the spade-foot (Scaphio-
podidae) it is inguinal. In the eight species of frog (Ranidae) considered, it is
pectoral. These eight are, southern meadow frog, gopher frog, carpenter
frog, green frog, mink frog, Heckscher's frog, southern bullfrog, and bullfrog.

CROSS EMBRACES

(PI. IV, Fig. 7)

In our study of the Salientia of Ithaca we were in the process several years
and had ample material. But in a trip to other parts one's material is care-
fully isolated to insure normal life history notes in the short period at hand.
In these trips little attempt or opportunity was given individuals for cross
embraces in the camp laboratory. Some of the records given were field
observations and casual notes.

General Discussion 29

Scaphiopus holbrookii cf X Hyla gratiosa 9

Bufo terrestris cf X Rana sphenocephala cf

Bufo terrestris cf X Rana sphenocephala 9

Bufo terrestris cf X Hyla gratiosa cf

(Rana septentrionalis cf X Rana clamitans 9 )

When ovulating is taking place in the same place for several species,
embraces might occur. For example, a Bufo terrestris male might leap at
Hyla femoralis, Hyla cinerea, Hyla gratiosa, Scaphiopus holbrookii or any
other if it approaches its size.

Some workers on the basis of few observations may think the males can
distinguish sexes but countless cases of males mating have been observed.
We did not record all we saw, but have in our notes the following species
where males amplexated :

Bufo quericicus Hyla squirella

Bufo terrestris Hyla versicolor

Acris gryllus Pseudacris ocularis

Hyla femoralis Rana sphenocephala

Or to make the case more pronounced males sometimes mate in a tier of
three : one female below with a male atop the male mating with her. We recall
such a case in Pseudacris ocularis, Bufo quercicus and Scaphiopus holbrookii.

Abnormal embraces with forms other than frogs were not sought or
observed.

DEPARTURES FROM CUSTOMARY AMPLEXATION AND
THEIR SIGNIFICANCE

(PL IV, Figs. 5, 6, 8)

In 1914 we remarked the emphasis laid by European workers on form of
embrace or amplexation in differentiating or diagnosing certain families or
genera. In Dr. Gill's comments on Boulenger's Tailless Batrachia he notes
that differences may in part be due to differences in size between males and
females and must vary.

Since the Ithaca paper we have seen the amplexation of almost every
species of the southeastern United States. In general with eastern Salientia
we can say that every species of Rana embraces pectoral-like; that every
Hylid (Acris, Pseudacris, Hyla), each Bufo (B. americanus, B. fowleri, B.
terrestris, B. quercicus), and the narrow-mouthed toad {Gastrophryne carolinen-
sis) are axillary or supra-axillary in fashion while the spade-foot is inguinal.
But every one of these may at times be varied from the normal.

Relative size, beginning, crest or waning of impulse or exhaustion of male
or presence of other males may dictate the form of amplexation.

In most of the species of Okefinokee when enough pairs were at hand
variations occurred.

Sometimes a male will seize a female in pre-axillary form (Gastrophryne) .
Or a small male may apparently have to be supra-axillary or pre-axillary in
front of forearms over the tympanum (Bufo terrestris). Or a male if it has

'^>;^>ii^^sSi

3© Frogs of the Okejinokee Swamp

amplexated axillary for several times over several days may be pre-axillary
with fore hands on lower throat or pectoral region (Acris), or on one side have
arm to middle of female's throat and other arm axillary (Hyla cinerea or
Bufo terrestris).

Axillary fashion in pectoral or inguinal species occurs. A Scaphiopus
male once held a female with one arm above and behind a foreleg and the
other just in front of a hind leg. Or a southern meadow frog, male, may
seize a female Rana clamitans axillary-like, not have arms long enough to
reach breast. Or an unusual pair of R. grylio might seem axillary like.

Pectoral amplexation might come in axillary forms when a male proves
bigger than the female (Hyla femor alts, Bufo terrestris, R. clamitans). Some-
times both arms go on to the breast or only one. The other arm might be
axillary like in embrace.

In toads and spade-foots ardor is very great. One toad we saw embracing
a floating rootstock. Any frog its size it will seize. In the same way the
spadefoot will seize southern meadow frogs, Florida tree frogs, southern toads
or similar-sized anurans.

In spite of the variations given in the three preceding paragraphs we have
never seen ovulation in any species where abnormal amplexation obtained
and in general most of the preceding records must be preceding ovulation or
where captives have long remained in embrace. Normally at ovulation in
the field and usually in laboratory amplexation is normal.

OVULATION

In our study of ovulation we sought to capture mated pairs afield. These
usually laid the night of their capture. There were a few exceptions to the
rule as we found for the forms of northeastern United States, but in general
all species follow it. It means that when the female appears at the breeding
place she is usually ready to ovulate. If mating males be not present or if a
female of a pair be separated from a mating male or if a female be caught in
a congress just before mating, she may lay in captivity without the attendant
male. We did not seek to see if such were possible for each species, but it is
without doubt the case. At random with no attempt to secure such results
we noted ovulation without attendant males in Acris gryllus, Hyla versicolor,
Hyla cinerea, Bufo terrestris, Rana clamitans, Rana sphenocephala and
Rana virgatipes.

In very early spring at least six forms begin to breed. Possibly Pseudacris
nigrita may lead or come after the others. Its eggs we have not found.
They are doubtless a loose small mass like those of P. triseriata or P. feriarum
farther north. The southern meadow frog Rana sphenocephala begins laying
its plinths from February to December; crest March to May. About the
same time, February 15 or 20 to August 17 (crest March to May) the gopher
frog Rana aesopus lays a larger plinth similar to that of the southern meadow
frog. March i to September i seems the period, (crest March to May) for
the long egg files of the southern toad Bufo terrestris. From March 10 to

Plate IV

Amplexation
Normal types.

1 . Pectoral, Ranidae, Rana pipiens, Ithaca, N. Y.

2. Inguinal, Scaphiopodidae, Scaphiopus holbrookii, Hilliard, Ha. August i6, 1922.

3. Axillary, Hylidae, Hyla versicolor, Ithaca, N. Y.

4. Axillary, Brevicipitidae, Gastrophryne carolinensis, Billy Id., Ga. June 29, 192 1.

Abnormal

5. Preaxillary, Gaslrophryne carolinensis, Billy Id., Ga. May 21, 1921.

6. Multiple amplexation, Pseudacris ocularis, The two upper are males, Billy Id.,

Ga. May 26, 1921.

7. Cross embrace of a male Bufo valUceps with a female Rana pipiens, Beeville,

Tex. March 27, 1925.

8. Two males in embrace, Hyla versicolor, Chesser Id., Ga. August 12, 1922.

Plate V
Normal embraces.
HyJa cinerea, Billy Id., Ga. Jul}' 7, 192 1. .\xillarv amplexation.
Hyla femoraUs, Billj^ Id., Ga. May 20, 1921. Axillary amplexation.
Hijla gratiosa, Coat -bet pond, Chesser Id., Ga. July 17, 192 1. Axillary amplex-
ation.

4. ////ksg»(VfZ/a, Anna's pond, Folkston, Ga. July 3, 1922. Axillary amplexation.

5. Acris gryllus. Startling Branch, Folkston, Ga. July 2, 1922. Axillary amplex-

ation.
Pseudacris ocularis, Billy Id., Ga May 26, 1921. Axillary amplexation.
Bufo quercicus, Billy Id., Ga. June 4, 192 1. Axillary amplexation.

8. Bufo terrestris, Bill}- Id., Ga. July 6, 1921. Axillary amplexation.

9. Gastrophryne carolinensis, Billy Id., Ga. May 21, 1921. Axillary amplexation.

10. Bufo foiiieri, Dinwiddie, Va. June i. 191 7. Axillary amplexation.

11. Rana clamitans, Billy Id., Ga. July 4, 1921. Pectoral amplexation.
Rana spheuocephala, Billy Id., Ga. Ma}- 22, 192 1. Pectoral amplexation.

Plate VI
Egg masses

1. Acris gnjlliis eggs, laid in camp, Billy Id., Ga. May i6, 1921.

2. Scaphiopus holbrookii eggs, Hilliard, Fla. August 17, 1922.

3 . Gastrophryne carolinensis eggs, Starling Branch, Folkston, Ga. .July

4. Pseudacris ocularis eggs, laid in camp, Chesser Id., Ga. July 17, 1921

General Discussion

31

August 17 (crest April to June) the solitary spadefoot may begin its laying of
bands (later cylinders). From March 18 to September i, (crest April to
June) the Florida tree frog (Hyla gratiosa) lays its single eggs. Thus we
have in early spring six forms — a small loose egg mass, two plinths, egg
strings, egg bands or cylinders, and single eggs. However different in form,
they are all alike in being submerged.

Our evidence points to six forms as beginning ovulation in April. The
southern tree frog, {Hyla squirella) lays single eggs in April to August
(crest April to June). The cricket-frog {Acris gryllus) apparently comes
next, April 15 to September i (crest May 10 to July 20) with its single eggs
(rarely in masses) . The cow bell frog ( Hyla cinerea) lays its surface films or
submerged masses from April 15 to August 15 (crest May 10 to July 15).
Soon after, April 20 or 25 to September i (crest May 20 to July 15) the pine
wood tree frog {Hyla femoralis) deposits its films on the surface. From
April 24 to August 21 (crest May 20 to July 20) the little chorus frog {Pseu-
dacris ocularis) emits single eggs on the bottom of ponds and on vegetation.
From the last of April to August 1 1 the carpenter frog {Rana virgatipes) lays a
plinth attached to sticks or stones like meadow frog masses, but the indi-
vidual eggs have no inner envelopes. Three of the six lay single submerged
eggs, one a plinth, one a surface film, and one sometimes a surface film, or
sometimes a submerged mass. Possibly Heckscher's frog {Rana heckscheri)
begins April i and stretches to August i (crest? May and June) but this
period is uncertain as is the conjectural mass. Is it a film hke R. grylio, R.
clamitans, or if in April, a submerged mass like R. virgatipes, R. sphenocephala,
or R. aesopus?

About May i four seem to start. From May i to August 1 1 (crest May
20- June 30) the common tree toad {Hyla versicolor) ought to deposit its
surface films. From May i to August 31, (crest May 20 to July 23), the
narrow-mouthed toad {Gastrophryne carolinensis) spawns clear marble-like
eggs in films on the surface, these eggs truncated on the top. From May i to
July 8 or later (crest May 20 to June 14) the Anderson tree toad might if in
Okefinokee region lay its single submerged eggs. One more goes from May i
onward to September i (crest May 23 to July 26) and it is the oak toad {Biifo
quercicus) which deposits short bars of eggs on the bottom of ponds.

From the middle of May to September i (crest May 24 to July 28) the
southern bullfrog {Rana grylio) has large films one by two and a half feet
across, much as does Rana catesbeiana. If this latter form be near the Oke-
finokee it doubtless lays the last of May or from June to August i when the
green frog {Rana clamitans) has been recorded laying its smaller surface
films. From May i onward we thus have remarked seven forms of which
five lay surface films and two submerged (one single eggs, one bars in different
forms) eggs.

Some of the spawning dates, order of ovulation and humidity (Table 12)
and temperature data (Table 13) appear in the following tables.

32

Frogs of the Okefinokee Swamp

Table 12

Species

Spawning Period

Pseudacris nigrita

Rana sphenocephala

Feb. to

Dec.

Rana aesopus

Feb. 15

Aug.

17

Bufo terrestris

Mar. I

Sept.

I

Scaphiopus holbrookii

Mar. 10

Aug.

17

Hyla gratiosa

Mar. 18

Sept.

I

Hyla squirella

Apr.

August

Acris gryllus

Apr. 15

Sept.

Hyla cinerea

Apr. IS

Aug.

15

Hyla femoralis

Apr. 20

Sept.

I

Pseudacris ocularis

Apr. 24

Aug.

21

Rana virgatipes

Apr. (last) Aug.

21

Rana heckscheri

(Apr. I?

Aug.

I?

Hyla versicolor

May I

Aug.

II

Gastrophryne carolinensis

May I

Aug.

31

(Hyla andersonii

May I

July

8

Bufo quercicus

May I

Sept.

I

Rana grylio

May 15

Sept.

I

(Rana catesbeiana

June

July

Rana clamitans

June 2

Aug.

10

Rana septentrionalis

June 25

Crest of Ovulation

Mar.

to

May

Mar.

May

Mar.

May

Apr.

June

Apr.

June

Apr.

June

May

10

July 20

May

10

July 15

May

20

July 15

May

20

July 20

May

July

May?

June?

May

20

June 20

May

20

July 23)

May

20

June 14)

May

23

July 26

May

24

July 28

June

20

July 15)

June

14

July 15
)

Table 13

Range

Average

Range Average Humidity

Pseudacris nigrita

Rana sphenocephala

50-74

65°

66-78

Rain

Rana aesopus

68-74

71

81-91

86

Inch or more of rain

Bufo terrestris

50-78

66

73-97

.90-2.60 inches

Scaphiopus holbrookii

Heavy rains

Hyla gratiosa

62-83

70

78-98

90

.91-2.40 inches

Hyla squirella

62-74

70

81-96

90

1. 42-1. 80 inches

Acris gryllus

58-73

69

73-97

89

Heavy rains

Hyla cinerea

60-74

75-91

.84-2.80 inches

Hyla femoralis

61-74

81-96

.48-3.47 inches

Pseudacris ocularis

70-79

74

Rains

Rana virgatipes

70-82

Rains

Hyla versicolor

68-74

83-96

91

1.48-4.06 inches

Gastrophryne carolinensis

66-77

72

78-97

89

T-5.00 inches

Bufo quercicus

64-74

79-96

T-5.31 inches

Rana grylio

68-78

85-96

Humid or rain

Rana clamitans

68-75

88-94

90

None-considerable

Rana heckscheri

Quite obviously the evidence of minima, etc. for Rana aesopus, Scaphio-
pus holbrookii, Pseudacris ocularis, Rana virgatipes and Rana heckscheri is
too scanty to be sure of their place in this table. In Okefinokee swamp
humidity is the great factor in precipitating ovulation.

General Discussion 33

THE EGGS
(Pis. VI, VII, VII, IX; Text Fig. i)

Incorporated with this consideration of our southeastern species is a
previous key to the eggs of our northeastern forms. The combination may be
termed "A key to the eggs of the Sahentia east of the Mississippi River."

Workers will discover the following species missing from the key because
the authors themselves have had no opportunity to secure both fresh and
preserved materials of these species:

Hyla andersonii — Eggs first found by T. Barbour and later by G. K.

and R. C. Noble.
Hyla c. evittata — Eggs found by M. K. Brady. Quite similar to H. cinerea.
Pseudacris ornata — Eggs not known. Described by J. K. Strecker.
Pseudacris nigrita — Eggs most certainly like Pseudacris triseriata, P.

feriarum, and P. septentrionalis.
Rana areolata— Eggs described by H. P. Wright and G. S. Myers.
Rana cantabrigensis — Eggs very similar to R. sylvatica.

The season of breeding for species in the north is marked both at the
beginning and at the end. Each species occupies four or five weeks except
Bufo americanus and Rana clamitans. The exceptions may require two or
three months for ovulation. In the southeastern states when once a species
has begun, its season of breeding may extend throughout the summer or
even into the early fall, dependent upon the high crests of precipitation.
These species, although of a swampy region, wait for the rains and in this
reliance on precipitation they suggest our desert species of Texas and Arizona.
Those which do not begin until June have at least eight to ten weeks of
ovulation. This minimum period for a species of the south is the maximum
period for a northern form. Species such as Rana sphenocephala, Bufo ter-
restris and Acris gryllus, which begin early in the season, breed during 25-30
weeks of the year, if not longer or from February to September or October.

The number of eggs in a complement may vary from 100 in the smallest
species, Pseudacris ocularis, to 20,000 in Rana catesbeiana, the largest form.
The range in the tree frogs (Hylidae) is from 100 (Pseudacris ocularis) to
2,084 (Hyla gratiosa); in the toads (Bufonidae) from 610 (Bufo quercicus)
to 8,000 (Bufo americanus); in the frogs (Ranidae) from 349 (Rana virgatipes)
to 20,000 (Rana catesbeiana). The complements of the narrow-mouthed toad
Gastrophryne carolinensis) and the spade-foot (Scaphiopus holbrookii) are,
respectively, 869 and 2,332.

The eggs of seven species, Hyla cinerea, H. femoralis, H. versicolor, Rana
catesbeiana, R. clamitans, R. grylio and Gastrophryne carolinensis float on the
surface of the water; the eggs of the other 17 species are submerged. In
northern or southeastern states no form with buoyant eggs lays before May
10. The II or 12 early breeders have submerged eggs. These are usually
with firm jelly envelopes except for Pseudacris triseriata and Scaphiopus hol-
brookii, which have the consistency intermediate between the firm jellies of
early breeders and the loose surface films of late breeders. One form. Gas-

34

Frogs of the Okefinokee Swamp

®

I {

r
®

8

©
1

®

II

n

Figure I. Individual Eggs. X 2.3

Bufo americanus
Bufo terrestris
Bufo fowleri
Bufo quericus
Scaphiopus holbrookii
Pseudacris ocularis
Pseudacris triseriata
(From Buffalo, N. Y.)

9-
10.

12.
13-
14.

15-
16.

Hyla crucifer
Acris gryllus
Hyla squirella
Hyla gratiosa
Hyla cinerea
Rana catesbeiana
Rana virgatipes
Rana grylio
Rana clamitans

Hyla femoralis
Hyla versicolor
Rana palustris
Rana pipiens
Rana sphenocephala
Rana aesopus
Rana sylvatica
Gastrophryne carolinensis

In No. 15 the vitellus was inadvertently drawn too large.

trophryne carolinensis, although it lays at the surface, has the most beautifully
distinct, firm eggs of all the species considered.

The difference in jelly consistency in an early spring laid egg and late
summer laid egg of the same species has not been studied for differences.
Nor have we fully determined whether heavier rainfall is needed for the tardy
breeders than for the early ones of a species. The eggs of Rana heckscheri,
and Pseudacris 7iigrita of the Okefinokee forms remain yet to be described
by someone.

The eggs were laid in camp and in the laboratory by mated pairs caught
in the field. Later the eggs in the field were determined by these original
checks. Occasionally the process of egg laying was observed in the field.
Two species, one of the north and one of the south, were identified by the
positive elimination of all the other resident forms.

General Discussion

35

The measurements and color descriptions are based on fresh eggs. Later,

these were checked with preserved material. The eggs with loose outer

envelope have the outer margin indicated by dots. In one species the vitelline

membrane is far separated from the vitellus and the space is indicated by

cross hatching. A summary of the egg characters of each species follows in

the accompanying key:

A. Eggs deposited singly,
a. Envelopes two

Outer envelope diameter 1.4-2.0 mm.; inner envelope diameter 1.2 to 1.6 mm;
vitellus diameter 0.8 to i.o mm; eggs brown above and cream below. Egg

complement, 942. Season, June 10 to August 21 (Fig. 10) Hyla squirella.

aa. Envelope single

b. Envelope 2.3 mm. or more.

c. Vitelline membrane far from vitellus, appearing as inner envelope 1.6 to
2.0 mm; outer envelope loose, glutinous, indefinite in outline 2.3 to 5 mm;
vitellus 1.0 to 1.8 mm. Egg complement, 2084. Season, June 10 to

August 21 (Fig. 11) Hyla gratiosn.

cc. No inner envelope or appearance as such, envelope firm, definite in outline
2.4 to 3.6 mm; vitellus 0.9 to i.o mm. Egg complement, 241. Season
April 15 or earlier to September i (Fig. 9) . . Acris gryllus.
bb. Envelope 1.2 to 2.0 mm.

c. Vitellus 0.6 to 0.8 mm. Egg complement, 100. Season, May 16 to August
21 (Fig. 6) . . . Pseudacris ocularis.
cc. Vitellus 0.9 to i.r mm. Egg complement, 800 to 1,000. Season, March 30

to May 15 (Fig. 8) Hyla crucijier.

AA. Eggs deposited in a mass.

a. Egg mass, a surface fUm.

b. Egg envelope outline always distinct, never lost in the mass; eggs firm and
distinct like glass marbles, making a fine mosaic; envelope a truncated sphere,
the flat surface above; envelope single 2.8 to 4.0 mm; vitellus 1.0 to 1.2 mm;
color black above and white below. Egg complement, 869. Season, May 21

to August 17 (Fig. 24) Gastrophryne carolinensis.

bb. Egg envelope outline indistinct, more or less merged in the jelly mass; jelly
glutinous; egg brown above, cream or yellow below.

c. Egg packets small, masses seldom if ever over 20 sq. in. (125 square centi-
meters), or 4 by 5 inches in diameter (10 by 12.5 cm.).
d. Inner envelope large 2.2 to 3. mm; outer envelope 3.2 to 5.0 mm;
vitellus 0.8 to 1.6 mm; Egg complement, 343 to 500. Season, May

19 to August 21 (Fig. 12) Hyla cinerea.

dd. Inner envelope small 1.4 to 2.0 mm; outer envelope 4 to 8 mm.

e. Packets small, seldom over 30 to 40 eggs; vitellus i.i to 1.2 mm.
Egg complement, 1,802. Season, May 10 to August 13 (Fig.

18) Hyla versicolor.

ee. Packets large, sometimes 100 to 125 eggs; vitellus 0.8 to 1.2 mm.
av. 0.95 mm. Egg complement, 768. Season May 16 to August

21 (Fig. 17) Hyla femoralis.

cc. Egg packets large, loose, glutinous films, 35 sq. in. to 675 sq. in. (218 to

3,721 sq. cm.).

d. Inner envelope absent; vitellus 1.2 to 1.7 mm; egg mass 144 to 675

sq. in. (900 to 3,721 sq. cm.) in area, or 12 by 25 inches (30 by6i cm.)

in diameter; egg masses amongst brush around the edge of ponds or

encircling Pottiederia-\ike vegetation in midpond. Egg complement,

10,000 to 20,000. Season, June i to July 10, (Fig. 13)

Rana catesbeiana.
dd. Inner envelope present, 2.8 to 4.0 mm; vitellus 1.4 to 2.0 mm.

e. Egg mass seldom i. sq. ft. (35 to 144 sq. in. or 218 to 900 cm) in
area or 5 by 7 to 12 inches in diameter; usually around edge of
ponds; inner envelope eUiptic, pyriform or circular, av. 3.05 mm;
vitellus 1.4 to 1.8 mm, mode 1.4 mm. av. 1.5 mm. Egg comple-
ment, 1,451 to 4,000. Season, May 2t, to August 21 (P'ig. 16) . . .

Rana cLamitans.
ee. Egg mass over i sq. ft. in area (144 to 288 sq. in. or 900 to 1,800
sq. cm.) or 12 by 12 inches to 12 by 24 inches in diameter; usually
in midpond; inner envelope av. 3.45 mm; vitellus 1.4 to 2.0 mm,
mode 1.8 mm, av. 1.7 mm. Egg complement, 8,000 to 15,000.
Season, May 24 to August 21 (Fig. 15) Rana grylio.

36 Frogs of the Okefinokee Swamp

aa. Egg mass submerged.

b. Eggs in files or bands.

c. Eggs laid in bands which soon become loose cylinders extending along
plant stems or grass blades; vitellus 1.4 to 2.0 mm; envelope 3.8 to 5.6 mm.
Egg complement, 2,332. Season, April 15 or earlier to August 17 (Fig.
5) Scaphiopus holbrookii.

cc. Eggs in files.

d. Files short (4 to 10 mm in length); 4 to 8 eggs in short bead-like chain
or bar or many such files radiating from one focus; vitellus 0.8 to i.o
mm; tube diameter 1.2 to 1.4 mm. Egg complement, 610, 766.

Season, June 4 to August 21 (Fig. 4) Bufo quercicus.

dd. Files long (several feet in length or often a meter or more long);
vitellus 1.0 to 1.4 mm; tube diameter 2.6 to 4.6 mm.
e. Inner tube absent; outer tube 2.8 to 3.4 mm; vitelli crowded in
the files; at first in a double row, later more spread out but still
crowded; 22 to 24 eggs in 30 mm (i 3/16 inches). Egg comple-
ment, 7,750. Season, April 15 or earlier to August 17 (Fig.

3) Bufo foivleri.

ee. Inner tube present.

f. 18 to 20 eggs in 30 mm (i 3/16 inches); partition apparent
between eggs; inner tube 1.6 to 2.2 mm; outer tube 3.4 to
4.0 mm. Egg complement, 4,000 to 8,000. Season, April 5

to July 25 (Fig. i) Bufo americanus.

ff. 7 to 8 eggs in 30 mm (i 3/16 inches); distinct space between
eggs; no partition apparent; tube inclined to be slightly
emarginate between the eggs; inner tube, 2.2 to 3.4 mm; outer
tube, 2.6 to 4.6 mm. Egg complement, 2,888. Season, April

15 or earlier to August 13 (Fig. 2) Bufo terrestris.

bb. Eggs|in|lumps.

c. Egg mass a firm regular cluster.

d. Egg mass a sphere 2^ to 4 in. (6.35 to 10 cm) in diameter, containing
2,000 to 3,000 eggs; outer envelope distinct.)

e. Eggs black above and white below; inner envelope apparently
absent, slightly evident under lens, 3.6 to 5.8 mm; outer envelope
5.2 to 9.4 mm; vitellus 1.8 to 2.4 mm. Egg complement, 2,000
to 3,000. Season, March 19 to May i (Fig. 23). . . . Raiia sylvatica.
ee. Eggs brown above and yellow below; inner envelope distinct 2.3
to 3.0 mm; outer envelope 3.6 to 5.0 mm; vitellus 1.6 to 1.9 mm.
Egg complement, 2,000 to 3,000. Season, April 6 to May 18

(Fig. 19) Rana palustris.

dd. Egg mass a plinth.

e. Without inner envelope, complement small, 349 to 474; eggs black
above and sulphur or primrose yellow below; eggs further apart
than in R. pipiens or R. spheiiocephala; outer envelope 4.9 to 6.9
mm; vitellus 1.5 to 1.8 mm. Egg complement, 349 to 474.

Season, June 21 to August 1 1 (Fig. 14) Rana virgatipes.

ee. With inner envelope, complement large, 1,000 to 5,000 or more;
eggs black above and white below.

f. Vitellus average 2.0 mm (range 1.8 to 2.4 mm); inner envelope
3.1 to 4.4 mm; outer envelope 4.4 to 6.0 mm, mode 5.2 mm,
av. 5.3 mm. Egg complement, 5,000 or more. Season, (?)

to August 17 (Fig. 22) Rana ae.^opus.

ff. Vitellus average 1.4-1.7 mm (range 1.3 to 2.0 mm); inner
envelope 2.3 to 3.2 mm; outer envelope 3.4 to 6.6 mm. Egg
complement, 1,000 to 5,000.
(g. Average outer envelope 5.0 or larger.

(h. Average outer envelope 5.1 mm. (range 4.2 to 6.0
mm, mode 5.0 mm.); vitellus 1.7 mm. average (1.4
mm-2.o mm). Egg complement, 3,500 to 4,500.
Season, March 30 to May 15 (Fig. 20) Raiia jripiens.
(hh. Average outer envelope 6.3 mm. (5.6-6.6 mm);
vitellus 1.4 mm. average (range 1.3 to 1.6 mm);
vitellus black or brown above, white or yellowish be-
low. Season, June 25 to July 25. Rana septentrionalis.
gg. Average outer envelope 3.8 mm (range 3.4 to 5.4 mrn, mode
4.0 mm). Egg complement, 1,054. Season, April 15 or
earlier to August 21 (Fig. 21) Rana sphenocephala.

General Discussion

37

Egg mass a loose irregular cluster.

d. Egg mass small, less than i in. (2.5 cm) in diameter. 20 to 100 eggs

in the mass; outer envelope merged ; the one envelope 5.0 to 7.8 mm,

rarely 3.0 mm; vitellus 0.9 to 1.2 mm. Egg complement, 500 to 800.

Season, March 19 to May i (Fig. 7) Pseudacris triseriata.

dd. Egg mass an irregular cyhnder i to 6 in. (2.5 to 15 cm) in length,
extending along plant stem or grass blade; envelope, single, 3.8 to
5.6 mm; vitellus 1.4 to 2.0 mm. Egg complement, 2,332. Season,

April 15 or earlier to August 17 (Fig. 5) Scaphiopus holbrookii.

(See also (c) under (b) Eggs in files or bands.)

EGG-LAYING PROCESS

In most of these southeastern frogs the actual ovulation and fertihzation
process has not been observed. Generally we would return after midnight
or later with a pair or more and had to sleep to conserve our energies for
daytime observations where and when congresses occurred. The opportunity
to observe this process was therefore often deliberately passed by. These
are points the resident naturalists can do better in any event. At this junc-
ture we will not enter into any discussion of the process, though we have
opinions, but in some cases not all the facts we want.

HATCHING PERIOD

Scaphiopus holbrookii

Gastrophryne carolinensis

Pseudacris ocularis

Hyla squirella

Hyla gratiosa

Hyla cinerea

Rana grylio

Bufo terrestris

Hyla femoralis

Bufo quercicus

Acris gryllus

1-2 to 6 days
ih-3 days
1 5- 1 1 days
1 5-2 days

2 days
2-3 days
2-32 days
2-4 days

3 days
3-3I days
3-4 days

Submerged masses
Surface films
Single submerged
Single submerged
Single submerged
Surface or submerged
Surface film
Submerged files
Surface film
Submerged bars
Single submerged

Hyla versicolor
Rana sphenocephala
Rana virgatipes
Rana clamitans
Rana aesopus
Rana catesbeiana
Rana septentrionalis

3-5 days
3-5 days
3-5 days
3-6 days
4-4^ days
4-5 days
5-13 days

Surface film
Submerged mass
Submerged mass
Surface film
Submerged mass
Surface film
Submerged mass.

From a glance it will be seen that the minimum period for each Okefinokee
species is from i to 4 days for hatching, or the maximum period for each
species from if to 6 days. This would imply temperatures of 6o°-8o° Fahren-
heit or higher if our table of 1914 (p. 19) be correct. I still beheve there is
much to be said for that table. However, if one observe the table herewith
given he will note that the shortest periods of hatching are not necessarily
the surface films. Of the first five species with lowest hatching periods, four
have submerged eggs. Strangely enough the single eggs in general have a
tendency to hatch quicker than the masses. Probably in Okefinokee Swamp
the egg complements of all species are under more uniform water conditions
in temperature than we have in the north.

38

Frogs of the Okefinokee Swamp

THE LARVAL PERIOD

According to the length of the larval period, the species of the Okefinokee
region fall into two major groups: those which transform during the season
in which the eggs are laid (14 or 1 5 species) and those which require one or more
years for their development (5 or 6 species). The data are as follows : (Table 14)

Table 14

Average

Vitelline

Larval periods

Egg

season

transf.
size

diameter
(mm.)

Larval period

Short larval period:

Scaphiopus holbrookii

Mar. 10

to Aug. 17

10 mm.

I . 4 to 2 . 0

14-30 days

Gastrophryne carolinensis

May I

Aug. 31

II

I .0 to I . 2

23 - 67

Bufo quercicus

May I

Sept. I

8

I . 2 to I . 4

33 - 44

Bufo terrestris

Mar. I

Sept. I

8

I .0 to 1.4

35 - 55

Hyla femoralis

Apr. 20

Sept. I

13

0 . 8 to 1 . 2

35 - 65

Pseudacris nigrita

40 - 60

Hyla squirella

Apr.

Aug.

12

0.8 to I .0

40 - 60

Hyla gratiosa

Mar. 18

Sept. I

20

1 . 0 to 1 . 8

41 - 6s

Hyla versicolor

May 10

Aug. 13

16

I.I to I . 2

45 - 60

Pseudacris ocularis

Apr. 24

Aug. 21

8

0.6 to 0. 8

45 - 70

Hyla andersonii

May I

July 8

12

I . 2 to I .4

49?- 74?

Acris gryllus

Apr. IS

Aug. IS

13

0 . 9 to I . 0

50 - go

Hyla cinerea

Apr. 15

Aug. IS

15

0 . 8 to 1 . 6

55 - 63

Rana sphenocephala

Feb.

Dec.

23

I . 3 to 2 . 0

67 - 86

Rana aesopus

Feb. 15

Aug. 17

27 to 5S

1 . 8 to 2 . 4

85 -106

Long larval period:

Rana virgatipes

Apr. (las

t) Aug. 21

27

I . S to 1 . 8

300 -36s

Rana clamitans

June 2

Aug. 10

25

1 . 4 to 1 . 8

300 -400

Rana septentrionalis

June 25

34

I . 3 to I . 6

36s -400

Rana heckscheri

Apr. I ?

Aug. I ?

35

( )

36s?-730?

Rana grylio

May 15

Sept. I

44

I . 4 to 2 . 0

36s -730?

Rana catesbeiana

June

July

53

I . 2 to I . 7

365^-730

The forms with long larval periods, namely carpenter frog, green frog,
mink frog, Heckscher's frog, southern bullfrog and common bullfrog are
generally mid-summer breeders, require 25-53 mm- for transformation size
(usually 30-53 mm.) and have uniform-sized vitelli (1.3-2.0 mm.) relatively
small for the transformation size required.

The forms in the short larval group fall into three groups: The first
assemblage of four — solitary spade-foot, narrow-mouthed toad, oak toad,
and southern toad — have small average transformation sizes 8- 11 mm., rela-
tively larger vitelli 1.0-2.0 mm., and may require a larval minima of 14 to
35 days. The second group comprehend nine forms, namely the tree frogs
(Hylidae) of the swamp. They have larger average transformation sizes
(12 to 20 mm) and actually smaller vitelli and larval minima of 35 to 55 days.
The average of the minimum sizes of the vitelli of the nine species is between
.8 and .9 mm. wherein in the first short larval group of four it is 1.15 mm.

Plate MI
Eggs of Hylas

1 . Hyla cinerea, eggs in a jar, Billy Id., Ga. May 19, 1922.

2. Hyla versicolor, packet in the field, Dinwiddle Va. June r, 191 7.

3. Hyla femoralis, packet, Billy Id., Ga. May 22, 1921.

4. Hyla cinerea, packet on edge of bonnet leaf, Billy Lake, Ga. May 2t„ 1921

5. Hyla squirella, eggs, stained, Chesser Id., Ga. July 4, 1922.

6. Hyla gratiosa, Chesser Id., Ga. July 16, 1921.

Plate VIII
Egg masses

1. Rana damitam egg mass, partly hatched in white clayey pool at Thompson's

Landing, St. Mary's River, Ga. July 9, 1922.

2. Bufo quercicus, single egg bar, on bottom of pond near Starling Branch, Folks-

ton, Ga. July 27, 1922.

3. Bufo quercicus, clustered egg bars, laid in camp, Billy Id., Ga. June 5, 192 1.

4. Rana grylio egg mass. Long Pond, Billy Id., Ga. Maj' 24, 192 1.

5. Bufo terrestris egg strings in field, Billy Id., Ga. June 6, 1921.

6. Bufo terrestris egg strings in camp, Billy Id., Ga. April 24, 1921.

3-

Plate IX
Rana egg masses
Rana septentrionalis, Hellgate ponds, Onekio, N. Y. July 14, 1923. x
Rana nrgatipes, Chesser Id., Ga. July 22, 1922. x i.o.
Rana sphenocephala, Billy Id., Ga. May 22, 1921. x 0.5.

4. Rana aesopus, Hilliard, Fla. August 17, 1922. x 0.45.

^ ^ ^

General Discussion

39

The last group in the short larval group comprise two species; the southern
meadow frog and the gopher frog. These have transformation sizes from
18-35 (average 23-30 mm.) larger vitelh if anything than the winter (long
larval) group and minimal larval periods of 67-85 days.

We wished to see if the larval period lengths in any way corresponded
with the relation of the egg diameters (vitelli) to the transformation sizes.
We divided the minimum transformation size by minimum vitellus size for
column one. For column two we used the maximum transformation size
and maximum vitellus. Column three is the mean of the two. The order of
this table (Table 15) accords quite closely with the table of larval period
lengths and there is surely a relationship. For example in the table a trans-
formed solitary spade-foot is in body length only six times as long as the
diameter of the egg vitellus from which it came. Or a transformed bullfrog is
thirty-five times as long as the diameter of the egg vitellus from which it came.

Table 15

I .

2.

3-

I. Scaphiopus holbrookii

6.07

6.00

6.03

2. Bufo quercicus

5-0

7-5

6.25

3. Bufo terrestris

6.5

7 ■ 5

7.00

4. Gastrophryne carolinensis

7.0

10. 0

8.5

5. Hyla andersonii

II .0

10.7

10.85

6. Pseudacris nigrita

? 10. 0

? 12.0

? II. 0

7. Hyla cinerea

14-3

10.6

12.45

8. Hyla femoralis

12.5

12.5

12. 5

9. Acris gryllus

10. 0

150

12.5

10. Hyla squirella

13-7

13.0

13-35

II. Hyla versicolor

II. 8

16.6

14.4

12. Rana aesopus

150

14.6

14.8

13. Rana sphenocephala

13.8

16.5

15-15

14. Pseudacris ocularis

16.6

13-7

15-15

15. Hyla gratiosa

18.0

12.7

15-35

16. Rana virgatipes

15-3

17.2

16.25

17. Rana clamitans

14.6

18.6

16.6

18. Rana septentrionalis

22.3

25-0

23-65

19. Rana heckscheri

20. Rana grylio

22.8

245

23-65

21. Rana catesbeiana

35-8

34-7

35-25

TADPOLES

(Pis. X, XI, XII, XIII, XIV)

The following table of tadpoles: — total length, greatest length of body, and
tail, greatest depth of body and transformation size reveals a correlated
ascending scale for all five characters. Usually the body length of a mature
tadpole will be sHghtly greater than the average transformation size, but in
several instances the same. Wherever it is smaller than the average trans-
formation size we have added the range of transformation size. Usually in

40

Frogs of the Okefinokee Swamp

most of the tadpoles the body depth averages from 1.6-2.5 times in the size of
transformation or body depth of tadpole in tail length from 2.5-4.5 times de-
pending on the species. The order of size for the tadpoles usually accords
with the order for size of adults with the exception of Bufo terrestris, Scaphio-
pus holhrookii, Acris gryllus, and corresponds very nicely with the order
given in table (Table 16) on transformation size.

Table 16

Average

Greatest

Greatest

Greatest

Greatest body

transf.

body

taU

length

length

size

depth

length

Pseudacris nigrita

? 23 . 0 mm

. ? 8.8 mm.

(8-12 mm.)

? 5 . 0 mm.

? 14. 2 mm.

Pseudacris ocularis

23.0

8.8

8

S-o

14.2

Bufo terrestris

24.0

9.8

8

5-0

14.2

Gastrophryne carolinensis 26. 4

10. 0

II (7-12)

4.8

16.4

Scaphiopus holbrookii

28.0

12.0

10

5°

16.0

Hyla squirella

32.0

12.0

12

6.5

20.0

Hyla femoralis

33°

10. 0

13 (10-15)

50

22.5

Hyla andersonii

350

12.0

12

5-5

23.0

Hyla cinerea

40.0

150

IS

6.0

25.0

Acris gryllus

42.2

130

13

6.6

29.2

Hyla versicolor

46.6

15.8

16 (13-20)

10. 0

30.8

Hyla gratiosa

50.0

19.0

20 (18-23)

12.0

31.0

Rana sphenocephala

62.5

25

23

15-0

37-5.

Rana clamitans

84.8

27.8

25

17.2

57-0

Rana aesopus

81.0

29.0

(27-35)

14.0

530

Rana virgatipes

92 .0

30.0

27

150

63.0

Rana septentrionalis

99 0

32.0

34 (29-40)

19-5

67.0

Rana heckscheri

950

41-5

35

17.0

53-5

Rana grylio

100. 0

35-6

44 (32-49)

20.0

64.4

Rana catesbeiana

142.0

450

53 (42-59)

38.0

97.0

A synopsis of the tadpoles of the Okefinokee species with a few hypotheti-
cal and other species (put in parentheses) follows :

Some may attempt to use this synopsis as a key. It may work and at
times it will not. To any worker who has had much field experience it is
apparent that tadpoles of closely related species like other animate things
are no respectors of man-made keys. Variations in individuals will occur.
This is a synopsis of mature larvae. Half grown larvae or larvae close to
transformation or almost transformed are often quite abnormal in the usual
characters used in larval descriptions. For example we raised Rana virgatipes
larvae from eggs but they did not attain a size large enough to get the labial
teeth well developed. We had to secure another series of mature larvae before
the description was completed.

Mouthparts. Boulenger (1898) labelled the labial teeth row from the upper
edge of upper labium inward toward the horny mandibles and from the lower
edge of the lower labium inward toward the mandible. The author (1914)
labelled the labial teeth "from the outer edge of the lip to the inner row nearest

General Discussion 41

the beak and the lower labial teeth from the row nearest the beak to the outer
one next the papillose border." Possibly when our North American forms
are described Boulenger's system may prove the better but we have kept
to the first for consistency and other reasons. In the Scaphiopodidae some-
times the small upper row of teeth in labial border is lost yet the variations
come most frequent in the small rows at either corner of the horny mandibles.
In the Ranidae the greatest loss comes also in this same region near the ends
of the horny beak. In the Hylidae, forms like Hyla crucifer, however, often
lose the third outer lower row, the goatee,
a . Mouth disk absent ; no labial teeth ; no papillae ; no honry beaks ; spiracle
median near anus; nostril within edge of mouth fold; eye on a canthus;
tadpoles (23-26.4 mm) small; black or grayish olive tadpoles with a

stripe on the middle of the tail musculature Brevicipitidae.

b. Tail sometimes with black tip; eyes plainly visible from ventral
aspect; inner face of upper labial edge with no black pointed escres-
cences ; lower mandibular prolongation light ; general coloration black
with purplish gray or hair brown dots; venter with white or yellowish
bands and large blotches; sides of body with same coloration; light
band at base of tail musculature prominent (even in alcohol). Egg
with truncate outer envelope, giving mass a mosaic appearance on
water's surface. Virginia — Florida — Texas, and up Mississippi

River to Indiana Gastrophryne carolinensis.

aa . Mouth disk present ; upper and lower labial teeth ; labial papillae ; horny
beaks present; nostril free of mouth; eyes dorsal close together; tail tip
rounded.

b. Spiracle sinistral; upper labial teeth not with two close rows on each
ridge; labial teeth 1/3, 2/2, 2/3, 2/4, 3/3, 3/4, 4/4 to 6/5; upper and
lower horny beaks, papillae on lower edge of labium absent or one
or two rows:
c. Anus median; spiracle lateral below the lateral axis (of tail mus-
culature projected forward) sometimes as much ventral as lateral;
upper tail crest extends on to the body to a vertical nearer hind
legs than the spiracle or only half way; viscera visible (in pre-
served specimens) through the skin of the belly,
d. Labial teeth 3/4 to 6/6; papillae extending completely
around the border of the labium except for a short toothed
median interval above (sometimes absent in one species);
papillary border not emarginate on each side; eyes nearer
mid-dorsal line than lateral outline, on lateral axis; tadpoles
24.5-65 mm. in length; spiracle a sht, very low on side, about
on the level of the mouth; in general very bronzy tadpoles;

myotomes of tail musculature well indicated

Scaphiopodidae.

e. Teeth 6/6, 5/6, 6/5, 5/5; inner papillae present; spiracle
equidistant between eye and base of hind legs or vent ;
eye i . 4-1 . 8 nearer tip of snout than to the spiracle,

42 Frogs of the Okefinokee Swamp

average 1.52; internasal space in interorbital space 1.28
-1.83, average 1.56; depth of body 1.74-2.5 in body
length, average 2.04; muscular part of tail in depth of
tail 1.45-2.5, average 1.98; last lower row of teeth
longer than horny beak. Last lower row of teeth i . 5
times in next to lowest row of teeth. Egg mass an ir-
regular cylinder, at first bandlike.

Massachusetts — Florida — Texas and Arkansas

Scaphiopus holbrookii.

dd . Labial teeth 2/3 ; papillae confined to the sides of the labium
(on lower half only in Bufo punctatus) upper and lower edges
toothed; papillary border on each side emarginate; eyes
sHghtly nearer lateral outline than mid dorsal Hne, above
lateral axis; spiracle small, a porelike opening; tadpoles 24-

28 mm. in length Bufonidae.

e. Papillae on upper and lower halves of lateral labial
margin ; some inner papillae.

f. Black or blackish tadpoles; in life intestine shows
through the skin of the belly; third lower row of
labial teeth long, i . 2-1 .6 in first lower row; third
lower row i. 0-1.5 ^^ horny beak; median space
between lateral parts of the 2nd row of upper labial
teeth small, contained 2 . 0-4 . o times in either lateral
half, not greater than lateral half,
g. Papillae plainly visible.

h. Tadpole to 24 mm; horny beak in upper
fringe 1.75, in first or second lower row
1.5; horny beak equal to or less than the
3rd lower row of teeth; median space be-
tween two parts of second upper row i . 4-
2 . 1 in either lateral part; third lower row
of teeth in first lower row i . 2-1 .4; two or
more rows of strong papillae from end of
upper fringe to end of third lower row,
sometimes 3-5 rows at side of labium;
lower loop of papillae far below level of
third row and with at least two rows of
papillae ; mouth in interorbital space i . o-
1.5, average i . 1 7 ; mouth larger than inter-
nasal space 1 . 2-1 . 6, average 1.36; depth
of tail in tail length 2 . 4-4 . 5, average 3.27;
eye nearer snout than spiracle i . o-i . 56,
average 1.2; nostril nearer eye than snout
1.1-1.8, average 1.48. Egg mass long
file, inner tube no partitions. North

Carolina — Florida — Louisiana

Bufo terrestris.

General Discussion

43

hh. Tadpole to 27 mm; horny beak in upper
fringe i . 2-1 . 5, in first or second lower row
of teeth I . i-i . 2 or i . 2-1 . 3 ; horny beak
greater than 3rd row of lower labial teeth;
median space i . 1 5-2 . o, 1-4, or i . 3-3 . o
in either lateral part; third lower row of
teeth 1 . 3-1 . 5 or 1 . 4-1 . 5 in first lower row ;
one row of weak papillae from upper fringe
to end of third lower row of teeth with a
few scattering papillae at the side of the
labium; lower loop with only 2 or 3 scat-
tering papillae beside the outer row of
weak papillae :

i . Mouth in interorbital distance .77-1. o,
average . 92 ; horny beak in upper fringe
1 . 2-1 .4; horny beak in first or second
row 1.1-1.2 times; third row in first
lower row i . 3-1 . 5 ; depth of tail in tail
length 1 . 25-2 . 7, average 1.97; spiracle
nearer eye than vent i .04-1 . 54, aver-
age 1.28; eye nearer snout than spiracle
1 .0-1 . 27, average i . 16; mouth larger
than internasal space i . 4-2 . 2, average
1.76; papillae of lower labial loop do
not extend under the end of the end of
the third row of labial teeth ; tail muscu-
lature in tail depth i . 26-2 .66, average
2 . 04 ; internasal space i . 2-1 . 6 in inter-
orbital distance, average 1.6; spiracle
1 .05-1 . 55 nearer eye than vent, aver-
age 1.28. Egg mass long file, partitions,
inner tube present. Eastern North
America, from Hudson Bay southwest

Bufo americanus.

ii. Mouth in interorbital distance 1.07-
-1.5; horny beak in upper fringe i . 4-
1.5; horny beak in first or second row
1. 2-1. 3; third lower row 1.3-1.6 in
first lower row; depth of tail in tail
length 2 . 88-3 . 83, average 3.33; spir-
acle nearer eye than vent 1.2 5- 1.7,
average 1.45; eye nearer snout than
spiracle i . o-i . 6, average i . 16 ; mouth
larger than internasal space i . i-i . 83,
average 1.47; papillae of lower labial
loop shghtly extend or do not at all

44 Frogs of the Okefinokee Swamp

extend under the end of the third lower
row; tail musculature in tail depth
1 . 6-2 .3, average 1.85; internasal space
1 . 5-2 . 16 in interorbital space, average
1.86; spiracle i . 2 5-1 . 7 nearer eye than
vent or base of hind legs, average i . 45.
Egg mass long file, no inner tube, some-
times two rows of eggs. Michigan — New
England — Florida — Texas Buf of owleri.
cc. Anus dextral; spiracle distinctly lateral on or near body axis;

d. Papillary border on side with an emargination ; tadpoles 50-
149 mm. in length; papillary fringe on upper labium extends
not at all inward beyond the end of the upper fringe of teeth
or only 1/8 to 1/16 of the length of the fringe; length of
horny beak in upper fringe of teeth i . o to 1.5 times; labial

teeth 2/3 or 3/3 or 3/4 or more Ranidae.

e. Labial teeth 2/3 occasionally 3/3 rarely 1/3; tadpoles
74-150 mm. in length; dorsal crest not extending to
vertical of the spiracle, but usually just ahead of the buds
of the hind legs; tail always elliptical not spathulate;
upper fringe of teeth equal to or slightly larger (never
1 . 5 times) the horny beak.

f . Tadpoles 74-84 mm. ; tadpoles usually transform the
same season they are born; transformation sizes 18
to 30 mm. average 24 mm. (except R. aesopus);
tadpoles (except in R. aesopus) not strongly pigment-
ed on belly, viscera plainly showing through skin (in
spirit specimens.) Egg globular or plinthlike be-
neath surface of water,
g . Body in tail 2 . 1 5-2 .85, average 2.6; depth of
tail in length of tail 2.6-3.5, average 3.0;
nostril i .0-1 . 5. nearer eye than snout, average
1.25; eye i. 0-1.3 nearer spiracle than snout,
average i . 1 2 ; median space between the second
upper labial row i . 0-2 . o times the length of
either lateral part of this row; third lower row
.33-. 66 shorter than the first or second rows;
tail covered with large prominent dark spots;
belly strongly pigmented, in spirits it looks
white; viscera not visible. South Carolina —
Florida — Louisiana. (Viosca) . Rana aesopus.
gg. Body in tail i .3-2 . 2, averages (i . 53, 16, 1.7);
depth of tail in length of tail 2 . 3-3 . 4, averages
2.7, 2.65,3.0; belly not strongly pigmented, in
spirits dark, viscera show through the skin.

General Discussion

45

h. Median space in second labial row 2-4
times the length of either lateral part;
third row of lower labial teeth .33-. 66
shorter than the first or second rows,
usually at least .50; eye nearer the snout
than spiracle or equidistant ; nostril nearer
eye than tip of snout; depth of tail in
length of tail 2 . 3-3 . 2, average 2.7; spi-
racle 1. 5- 1. 8 nearer eye than snout,
average 1.63. Hudson Bay to Louisiana
and Eastern States .... Rana palustris.
hh. Median space .5-1.0 or i. 0-1.5 times
either lateral part of the second upper row;
third row of labial teeth .22 or .285-. 33
shorter than the first lower row; depth of
tail in length of tail 2 . 0-2 . 8 or 2 . 7-3 . 4.
i . Median space of second upper row . 5-
i.o times either lateral part; third
lower row of teeth .285-. 33 shorter
than the first lower row; nostril to snout
equal to nostril to eye; eye 1. 15- 1.3
nearer tip of snout than spiracle, aver-
age 1.2; body length in tail length
1. 3 5-1. 66, average 1.5; mouth .9-
1 . 6 larger than internasal space ; depth
of tail in length of tail 2 . 0-2 .88, aver-
age 2.65; tail crest usually with quite
prominent dark spots; greatest length
of tadpole 74 mm; spiracle 1.1-1.86
nearer eye than snout, average i . 46.
Southeastern States to Louisiana . .

Rana sphenocephala.

ii. Median space i. 0-1.5 times either
lateral part ; third lower row . 2 2 short-
er than the first lower row; nostril i . i-
1 . 5 nearer the eye than snout ; eye
nearer spiracle than snout 1.1-1.3;
body length in tail length i . 3-2 . o,
average 1.7; mouth larger than inter-
nasal space; depth of tail in length of
tail 2.7-3.4, average 3.0; tail crest
usually translucent with fine spots or
pencillings; greatest length of tadpole
84 mm; spiracle 1.4- 1.8 6 nearer eye
than snout, average i . 59. North

46 Frogs of the Okefinokee Swamp

America east of Sierra Nevada south-
ward into Mexico . . Rana pipiens.
ff. Tadpoles 84 to 142 mm; tadpoles usually winter
over at least one season; transformation sizes 28 to
59 mm. (except R. virgatipes 25 to 35 mm., possibly
R. onca) ; tadpoles usually with strongly pigmented
bellies, viscera not plainly showing through the skin
(in spirit specimens).

g . Tadpole with prominent continuous black crest
margins and a black musculature band; belly
bluish; tadpoles to 95 mm.; young tadpoles
black with transverse yellowish band on the
body; spiracle .86-1.2 nearer vent or base of
hind legs than snout, average i . o; spiracle . 85-
1 . 2 nearer eye than base of hind legs or vent,
i. e., usually equidistant; eye equidistant from
spiracle and tip of snout ; muciferous crypts very
distinct; spiracle below lateral axis; tail tip
acuminate; second upper labial row in upper
fringe 1/3 to 1/4; upper fringe distinctly
greater than horny beak ; median space between
two parts of second upper labial row i-i 1/2
of either lateral part; third labial row equal to
horny beak; third lower labial row longer than
single row of lower papillae; third lower labial
row 1/4 to 1/5 shorter than ist lower row. Eggs
unknown. South Carolina — Mississippi . . .

Rana heckscheri

gg. Tadpole without black crest margins or lateral

band; belly white, cartridge buff or yellow to

maize yellow; no transverse yellow band in

young tadpoles ; spiracle nearer vent than snout

1. 1 -1. 8; spiracle to eye rarely less than 1.25

greater than spiracle to vent; eye nearer tip of

snout than spiracle i .-i . 4; second upper labial

row in upper fringe 1/4 to 1/15; upper fringe

equal to or slightly greater than horny beak;

median space in either lateral part i 1/2 to 11 ;

third lower labial row much less (about i 1/2)

than horny beak; third lower row much shorter

or equal to single row of lower labial papillae.

h. Tadpoles to 140 mm; eye well above

lateral axis; muciferous crypts indistinct;

spiracle just below lateral axis; spiracle

1 . 06-1 . 44 nearer base of hind legs or vent

than tip of snout, average i .25; depth of

Plate X

Mature tadpoles, x i.o

• r . Hyla andersonii (two tadpoles ), Lakehurst, N. J. June 28, 1923.

2. Hyla cinerea. Chesser Id., Ga. August 12, 1922.

3. Hyla crucifer, Ithaca, N. Y.

4. H yla femoralis (two tadpoles) Anna's pond, Folkston, Ga. June 27, 1922

5. Hyla gratiosa, Petty pond, Folkston, Ga. July 31, 1922.

6. Hyla gratiosa, young tadpoles. Trader's Hill, Ga. June 24, 1922.

7. Hyla squirella (two tadpoles) Anna's pond, Folkston, Ga. June 27, 1922.

8. Pseudacris ocularis, Saddlebag pond. Trail Ridge, Folkston, Ga. June 30, 1922.

9. Hyla versicolor, Camp Pinckney, Ga. June 22, 1922.
10. Hyla versicolor, Ithaca, N. Y.

ir. .4cm gryllus. Board Pile Pt.. Ga. July 14, 1922.

12. Sca-phio-pus holbrookii, C&m^iYmckneY, Gsi. June 22, 1922.

13. Bufo quercicus. Chesser Id., Ga. August 10, 1922.

14. Gnsfrophryne carolinensis. Trader's Hill, Ga. June 24, 1922.

'"^t^

/

lO

12

Plate XI
Rana tadpoles. Mature, x i.o

1 . Rana aesopus, Chesser Id., Ga. August lo, 1922.

2. Rana aesopus, Petty pond, Folkston, Ga. July 29, 1922.

3. Rana clamitans, Near Spanish Creek, Folkston, Ga. July 17, 1922.

4. Rana clamitans, Ithaca, N. Y.

5. Rana grylio, Billy Id., Ga. May 11, 1921.

6. Rana grylio, Billy Id., Ga. August 4, 1922.

7. Rana heckscheri, Alligator Swamp, Callahan, Fla. July 20, 1922.

8. Rana heckscheri. Young tadpoles, Thompson's Landing, St. Mary's River, Fla.

July 17, 1922.

9. Rana sphenocephala, Chesser Id., Ga. July 25, 1922.

10. Rana sphenocephala, Anna's pond, Folkston, Ga. July 10, 1922.

1 1 . Rana septentrionalis, Onekio, N. Y. July 4, 1923-

12. Rana pipiens, Ithaca, N. Y.

13. Rana virgatipes, Lakehurst, N. Y. July 4, 192.V

14. Rana pahistris, Ithaca, N. Y.

lO

12

General Discussion 47

tail in tail length of tail 2.4-3.5, average
2.8; tail tip obtuse ; second upper row in
upper fringe 1/4 to 2/7; median space in
second upper row i 1/2 in either lateral
part; third lower row in first lower row
1/4-1/ 5 shorter; teeth 2/3 rarely 3/3.
Transformation size 43-59 mm. Egg
mass, a surface film. North America east of

Rockies Rana catesheiana.

hh. Tadpoles to 84-100 mm.; eye on or just
above lateral axis; tail tip acute or acu-
minate (rounded in R. onca) teeth 2/3;
second upper row in upper fringe 1/6 to
1/15: median space in second upper row
2.5-11 in either lateral part; third lower
row in first lower row 1/2-1/3 shorter,
i . Depth of tail in length of tail i . 45-1 • 8,
average 1.7; tail tip acuminate; dorsal
crest equal or less than tail muscula-
ture; muciferous crypts indistinct;
spiracle 1.08- 1.44 nearer vent than
snout; mouth in interorbital distance
1 . 5-2 .37, average 1.94; internasal
space in interorbital distance i . 8-2 . 6,
average 2.16; 2nd upper row 1/6 to
1/8 of the upper fringe; median space
of 2nd upper row 2 1/2 to 4 1/2 times
either lateral row ; third lower row i . 5
less than horny beak, much shorter
than single row of lower labial papillae
and I /3 shorter than first lower row of
teeth; ist row of lower teeth equal to
horny beak. Transformation size 32
or 37-48 mm.; Egg mass, a surface
film. Georgia — Florida — Louisiana . .

Rana grylio.

ii . Depth of tail in length of tail 2.5-4.7
averages 3.1-3.87; tail tip acute;
dorsal crest less than tail musculature ;
muciferous crypts distinct; spiracle
1. 07-1. 08 nearer vent than snout;
mouth in interorbital distance i . 3-1 . 8,
average 1.5; internasal space in inter-
orbital space 1.25-2.0, averages 1.6-
1.75; second upper row of teeth equal
to or greater than horny beak;

Frogs of the Okefinokee Swamp

Tadpoles to 99 mm.; transforma-
tion sizes at 29-38 mm; depth of
tail in length of tail 3 . 2-4.7, aver-
ages 3.87; spiracle just touches
lateral axis; eye just above lateral
axis; spiracle i . 06-1 . 38 nearer eye
than base of hind legs or vent, av-
erage 1.24; spiracle i . 25-1 . 6 near-
er eye than vent, 1.45; mouth in
interorbital distance i . 3-1 . 75, av-
erage 1.55; width of body in its own
length 1 . 3-2 . 1, average 1.56;
third lower labial row of teeth 1.25
less than horny beak, about equal
to single row of lower labial papil-
lae, 1/3 shorter than ist lower row;
sometimes a row of inner papillae
below the 3rd lower row of teeth;
median space in second upper
labial row 3 . 5-4 . 5 times either
lateral portion; second upper row
1/6 to 2/15 of the upper fringe;
belly straw yellow, colonial buff or
deep colonial buff; tail with round
cartridge buff or pinkish cinnamon
spots; no black line in dorsal crest
as in Rana grylio or R. virgatipes.
Eggs in a compact submerged mass.
Hudson Bay — Minnestoa — New

York — New England

Rana septentrionalis.

Tadpoles to 92 mm,; transforma-
tion sizes at 25-38 mm. depth of
tail in length of tail 2.5-3.7, spi-
racle just below lateral axis; eye on
lateral axis; third lower labial row
of teeth 1 . 5-1 .25 less than horny
beak, much shorter than single row
of lower labial papillae, almost
1/2 shorter than ist lower row;
median space of second upper row
6-1 1 times the length of either
lateral portion; second upper row
2/15 to i/i 5 of the upper fringe,
k. Spiracle nearer vent than
snout, 1. 3 5-1. 8; mouth 1.3-

General Discussion 49

1 . 8 times in interorbital dis-
tance, average 1.5; width of
body in its own length 1.25-
1.7, average 1.47 belly deep
cream color ; tail green mottled
with brown and covered with
fine yellow spots.
Egg mass, surface film.
Canada — Louisiana, Florida,
New England Rana clamitans.
kk. Spiracle nearer vent than
snout 1 .07-1 .45 averages
1. 23-1. 3; mouth 1. 0-1.3 7 in
interorbital distance, average
1 . 12.

1. Teeth 2/3 or 1/3; four to
six rows of inner papillae
from end of upper fringe to
end of lower labial row; a
row of heavy inner papil-
lae below the third lower
row of teeth ; spiracle near-
er eye than vent 1.42-
1.82, average i . 62 ; nostril
nearer eye than snout ; i . o
-1 . 42, average 1.2; width
of body in body length
I. 45-1. 86, average 1.6;
tadpoles large (92 mm.);
second upper row 2/15 or
1/15 of the upper fringe or
second upper row absent;
belly pale chalcedony yel-
low, sulphur yellow, vin-
aceous, pale grayish vin-
aceous or vinaceous buff;
tail: upper tail crest with a
black line or row of large
black spots, more promi-
nent than in R. grylio;
middle of musculature with
another black line; tail dark
with pale chalcedony yellow
spots. Transformation size
25-35 mm. Eggs, a sub-
merged mass .

50 Frogs of the Okefinokee Swamp

New Jersey to Georgia
. . . Rana virgatipes.
dd . Papillary border on side of labium without an emargination ;
tadpoles 23-50 mm. in length; labial teeth 2/3 or 2/2 . .

Hylidae.

e. Labial teeth 2/2; eye dorsal just inside the lateral out-
line in dorsal aspect (more like Ranids) eye i. 0-1.66
(av. 1.22) nearer tip of snout than spiracle; depth of
tail in length of tail 3.25-5.0, average 4.0; suborbital
region oblique not vertical; spiracle to eye usually equal
distance from spiracle to vent or base of hind legs;
spiracle plainly showing from dorsum; spiracular tube
in life stands out at an angle from the body and opening is
apart from the body proper; tail tip conspicuously black
(at times lost); papillary border does not extend above
the end of the upper fringe (Uke Rana) ; length of horny
beak in upper fringe of teeth i. 2-1. 3 5 times; Eggs
single, rarely a mass.
New York — Florida — Texas, up Mississippi Valley to

Canadian Northwest Acris.

ee. Labial teeth usually 2/3 (rarely 2/2) ; eye lateral, visible
from ventral as well as the dorsal aspect; eye i . o-i . 75
nearer the spiracle than tip of snout; depth of tail in
length of tail 1.6-4.4, average 2.1-3.55; suborbital
region vertical; spiracular tube in life parallel with body
and opening at inner edge closely connected with or

near to body proper Hylidae exclusive of Acris.

f . Labial teeth 2/2:

g. Teeth occasionally 2/2; tadpole to 33 mm.
spiracle 2 . 0-2 . 6 nearer base of hind legs than
tip of snout, average 2.16; nostril to eye 2 . o in
nostril to snout; tail crests clear, heavily pig-
mented with purplish black blotches on the
outer edge ; no prominent dark lateral band with
a clear light band below; upper fringe not per-
ceptibly angulate in the middle; two rows of
papillae on lower labial border below second
lower row of teeth; median space between sec-
ond upper rows of teeth 2-3 in either lateral
row; ends of second lateral row extending be-
yond end of upper fringe for 1/4 to 1/6 of the
length of either upper lateral row; horny -beak
1 . 75-2 . 25 in length from one end of lateral row
to end of the other lateral row. Eggs, single, sub-
merged. Manitoba — New Brunswick, South
Carolina, Louisiana Hyla crucifer.

General Discussion 51

Labial teeth 2/3:

g Third row of labial teeth short, shorter than
horny beak or . 20- . 40 of the first lower row in
length; upper fringe slightly or not angulate at
all; no flagellum ordinarily present; tadpoles
23-50 mm.; hght papillary development, lower
labial corner not with 3 or 4 strong rows of
papillae ; one or two rows of papillae beiow third
lower row of teeth; the papillae extend above
and beyond the end of the upper fringe for
about . 1 4-. 285 of the length of the upper fringe,
h. Tadpoles 23-33 mm.; eye equidistant be-
tween spiracle and tip of snout; spiracle
1 . 4-2 . 6 nearer vent or base of hind legs
than tip of snout; spiracle i .0-1 .6 nearer
eye than vent; papillae extend above the
fringe for .16- .25 of the length of the
fringe.

i. Musculature with no distinct brown
lateral hand with light area helow ; crests
usually heavily pigmented with purpl-
ish black blotches on outer rim ; nostril
to eye in nostril to snout 2.0; depth of
tail in tail length 2.4-3.15, average
2.7; spiracle 2 . 0-2 . 6 nearer vent than
snout, average 2.16; no papillae helow
third lower lahial row of teeth, thus ap-
pearing as a goatee; median space be-
tween second upper labial row 2-3 in
either lateral portion; horny beak in
upper fringe 1.75-2.25. Eggs single,

submerged Hyla crucifer.

ii. Musculature with a distinct hrown la-
teral hand with light area helow; crests
usually clear with fine scattered fleck-
ings, sometimes with fleckings gathered
nearer outer rim; one or two rows of
papillae below the third lower labial
row of teeth ; nostril to eye in nostril to
snout 1.25-2.0; depth of tail in tail
length 2.9-4.4, average 3.4, 3-55;
spiracle i . 1 5-2 . i nearer vent than
snout, average 1.5- 1.87 5; median
space between second upper labial row
2 1/2 or 3-7 in either lateral portion;
horny beak in upper fringe 1.5-2. o.

52 Frogs of the Okefinokee Swamp

j . Dorsal crest to vertical of spiracle;
spiracle i . o-i . 66 (average 1.33)
nearer eye than vent ; spiracle i . 1 5
-1.95 (average 1.5) nearer vent
than snout; mouth i . o-i . 25 larger
than internasal space; two rows of
papillae below third lower row of
teeth; third labial lower row .25-
.33 of the first lower row; first and
second lower labial rows i . 25-1 . 6
greater than the horny beak; horny
beak in upper fringe 1.5 -1.75;
median space between second row
of upper lower row 2 1/2 or 3-7 in

either lateral portion

Pseudacris (Raleigh.)

jj . Dorsal crest to vertical midway be-
tween spiracle and eye; spiracle
equidistant between eye and vent ;
spiracle i . 7-2 . i (average 1.875)
nearer vent than snout ; mouth and
internasal space equal; one row of
papillae below the third lower row
of teeth; third lower labial row of
teeth .33 of the first lower row;
dorsum of body in life with definite
scattered black spots; musculature
with three bands, apricot buff
(light) chestnut brown( dark) mar-
tins yellow (light); first and second
lower labial rows 2 . o greater than
the horny beak; horny beak in
upper fringe 2.0; median space be-
tween second row of upper labial
teeth 3-4 in either lateral portion.
Eggs single.

South Carolina — Florida — Louis-
iana, (Viosca) Pseudacris ocularis.
hh. Tadpoles 35-50 mm.; eye i .0-1 . 75 nearer
spiracle than snout ; spiracle i . o-i . 6 near-
er vent than the tip of the snout; spiracle
1 .25-2 . 5 nearer eye than vent or base of
hind legs; papillae extend above the upper
fringe for .14-. 285 of the length of the
fringe.

General Discussion 53

i. Tadpoles 50 mm. in length; body in
tail 2.3-3.25, average 2.5; depth of
body in width of body .83-1.0, aver-
age .9; depth of tail 10-14 mm.;
beautiful green tadpoles; young tad-
poles with a black saddle spot on the
back of the musculature near its base
and with a Ught Hne from eye to tail;
one row of papillae below lower third
labial row; papillae extending above
upper fringe for . 25-. 285 of the fringe's
length; dorsal crest extending to a
vertical halfway between eye and
spiracle. Eggs single, submerged.
South CaroHna— Florida— Louisiana.

(Viosca) Hyla gratiosa.

ii . Tadpoles 35-45 mm. ; body in tail i . i-
-2 . o, average 1.6; depth of body in
width of body i . o-i . 8 ; depth of tail 5-
9 mm. ; no black saddle spot in young
tadpoles.

(j. Tadpole small (35 mm.); dorsal
crest extending to vertical halfway
between spiracle and the base of
the hind legs; depth of tail in tail
length 2.5-3.5; average 3.0; nostril
to eye i . 2-2 . i in nostril to snout ;
mouth in interorbital space 1.33-
2.6; internasal space in interorbital
distance i . 33-2 . 2 ; eye just touches
lateral axis or is below it; horny
beak in upper fringe, 1.5-1.7;
papillae extending beyond the end
of the upper fringe . 25-. 285 of the
length of the upper fringe; two
rows of papillae below third lower
third lower labial row; median
space between second upper labial
row 1 . 2 5-2 . o in either lateral
portion; third lower labial row . 20-
. 22 of the first lower row; first row
of lower labial teeth i . o-i . 5 times
the horny beak. Eggs strewn in water
amongst sphagnum (Noble &Noble . )
New Jersey — South Carolina .

Hyla andersonii.

Frogs of the Okefinokee Swamp

jj . Tadpole medium (40 and 45 mm.)
Dorsal crest extends ahead of
spiracle or to eye; depth of tail in
tail length i . 5-3 . 2, average 2.75;
nostril to eye i . o-i . 7 in nostril to
snout; mouth in interorbital space
1.4-2.0; internasal in interorbital
space 1 . 2 5-2 . o ; eye on lateral axis ;
papillae extending beyond end of
upper fringe i . 4-2 . 5 of the length
of the upper fringe ; median space in
second upper labial row 3-5 in
either lateral portion; third labial
row . 25-. 40 of the first labial lower
row; first row of lower labial teeth
1.0^1.3 greater than horny beak,
k. Dorsal crest to the vertical
halfway between spiracle and
the eye ; depth of body in body
length 1 . 7-2 . 5 ; musculature
of tail in depth of tail i . 7 5-2 . 4
average 1.9; spiracle i . 4-2 . 3
nearer eye than vent; mouth
I . o-i . 4 larger than internasal
space, average 1.25; two rows
of papillae below the third
lower row of labial teeth; pa-
pillae extend beyond the end
of the upper fringe . 22-. 25 of
the length of the fringe ; horny
beak in upper fringe 2 . 0-2 . 3 ;
third labial lower row . 25-. 40
the length of the first lower
row.

Eggs surface or submerged ir-
regular mass.
Virginia — Florida — Texas to

Illinois Hyla cinerea.

gg. Third row of labial teeth long, longer than
horny beak, or . 75-1 . 00 of the first lower row
in length ; upper fringe very angulate in middle ;
flagellum on tail; tadpoles 32-50 mm.; heavy
papillary development, lower labial corner with
3 or 4 rows of papillae; two more or less com-
plete rows of papillae below third row of teeth
(except in Hyla arenicolor) ; papillae extend

Plate XII

Tadpole mouthparts ( Hyla, Acris, Pseudacris, Bufo)

1 . Hyla crucifer.

2 . Hyla gratiosa.

3 . Hyla andersonii

4 . Hyla cinerea.

5 . Pseudacris feriar u m .

6. Acris gryllus.

7 . Bitfo fouieri.

8 . Pseudacris ocularis

9 . Hyla squirella.

10. Bufo americanus.

1 1 . Bufo terrestris.
1 2 . Hyla versicolor
1 3 . Hyla femoralis.

Plate XIII
Tadpole mouthparts (Rana)

1 . Rana catesbeiana.

2 . Rana sphenocephala.

3. Rana palustris

4 . Rana aesopus.

5. Rana pipicns.

Plate XIV
Tadpole mouthparts (Rana, Gastrophryne, Scaphiopus)
Nana heckscheri.
Yg. Rana heckscheri.

3 . Rana septentrionalis.

4. Rana virgatipes.

5 . Rana grylio.

6. Gastrophryne carolinensis.
Rana clamilans.
Scaphiopus holhrookii.

==-?M

6

General Discussion 55

above and beyond the end of the upper fringe
for about . 30- . 40 the length of the upper fringe,
h. Third lower labial row .80-1.00 of the
length of the first lower row; dorsal crest
extends to the vertical halfway from hind
legs to spiracle, to spiracle or halfway from
spiracle to eye ; dorsal crest equal to, great-
er or less than depth of tail musculature;
tadpoles 36-50 mm.; red may be present
in the tail; tail crest distinctly or more or
less clear of spots next the musculature;
tail heavily blotched with dark blotches or
spots.

i. Medium space between lateral upper
rows 5.0-10.0 times in either lateral
row; spiracle i . 44-2 • 5 nearer eye than
vent; width of body in its own length
1.6-2. 1 ; eye i. 0-1.7 nearer spiracle
than tip of snout ; tail sometimes suffus-
ed with coral red, coral pink, or "red-
dish" or "orange."

j . Medium space between lateral
upper rows of teeth contained 6 . o-
1 0.0 in either lateral row; ist and
2nd lower rows of teeth i . 4-1 . 6
greater than horny beak; mouth
equal to internasal space ; depth of
tail in length of tail 1.6-2.75, av-
erage 2.25; muscular part of tail in
depth of tail 1.8-2.3, average 2.1;
depth of body 1.33-2.2 in body
length, average 1.68; dorsal crest
usually equal to or greater than
musculature depth; center of belly
soHd sulphur yellow; tail 3-5
handed] Hght lateral band bounded
below and above by a brown band;
flagellum clear of pigment; body
olivaceous black.
Eggs a surface film.
North Carolina— Florida— Texas

Hyla femoralis.

ii. Medium space between lateral upper
rows contained 3.25-5.0 times in
either lateral row; spiracle 1.12-1.5
nearer eye than vent; eye about equi-

56 Frogs of the Okefinokee Swamp

distant between spiracle and tip of
snout; internasal space in mouth . 7-
1 .0; dorsal crest extends to vertical of
spiracle or halfway between eye and
spiracle ; dorsal crest equal to or greater
than musculature depth; muscular part
of tail in depth of tail i . 72-1 . 9, aver-
age 1.8; depth of tail in length of tail
3.1-4.0; width of body in its own
length 1 . 3-1 . 7 ; no lateral hands in
tail; tail more or less scarlet or orange
Vermillion with Mack blotches more prom-
inent near the margins of the crests.
Body olive green; belly conspicuously
white or very light cream. Eggs a
surface film.

Minnesota — Texas — Maine — Flor-
ida Hyla versicolor.

hh. Third labial lower row . 75 of the length of
the first row; dorsal crest extends to the
vertical of the posterior edge of the eye;
dorsal crest usually less than depth of the
musculature; tadpoles to 32 mm.; width of
body in its own length 1.7-2.2, average
1.875; depth of tail in length of tail 2 . 2-
3.3, average 2.8; third lower row of teeth
not equal to first lower row; median space
between lateral upper rows contained 3.25
-5 . o times in length of either lateral row ;
papillae extend above and beyond the ends
of the upper fringe for .3-. 33 of the upper
fringe ; horny beak in upper fringe i . 8-2 . o ;
no bands nor red in tail; tail crest clear,
uniformly sprinkled with distinct black dots:
body greenish (like H. cinerea or Hyla grati-
osa) ; belly testaceous, or chalcedony yellow.
Eggs single, submerged.
Texas — Indiana; Florida to Virginia.

Hyla squirella.

TRANSFORMATION PERIOD

(Pis. XV, XVI)

Transformation for some species in Okefinokee may begin by April i or
earHer, but it occurs principally from May to August, especially from June
10- July 27. Thereafter to October i or 15 or November i, stragglers may
transform. The approximate order of transformation is given in the following
condensed table. (Table 17)

General Discussion

57

Table 17

Spawing
order

Larval
period

Transfc

)rmation dates

Average

Range

Rana clamitans

20

Over

April or May

Apr. I to Sept.

Rana grylio

18

I

June I

Apr. 24 to July ig

Rana heckscheri

13

winter

June 10

May to July 15

Rana virgatipes

12

June 14

May 15 to Aug. 15

(Rana catesbeiana

19

June

May to Oct. i)

(Rana septentrionalis

21

June

June or earlier to

September)

Pseudacris nigrita

I

Short

(Eariy)

to July I.

Rana sphenocephala

2

larval

June 21

-Apr. I to Oct. I

Rana aesopus

3

period

.

May i(?) to Nov. i

Bufo terrestris

4

in the

June 15

Apr. 27 to Oct. I

Scaphiopus holbrookii

5

year eggs
are laid.

Apr. 15 to Oct. I

Acris gryllus

8

" "

July 14

May 29 to Oct. i

Gastrophryne carolinensis

IS

" "

July 13

June 12 to Oct. 15

Hyla femoralis

10

" "

July 17

June 16 to Oct. i

Hyla squirella

7

" "

July 3

June 26 to Oct. i

Hyla versicolor

14

" "

June 27 to Aug. i

Hyla andersonii

16

" "

July 19

June 29 to Sept.

Pseudacris ocularis

II

" "

July 15

June 30 to

Hyla gratiosa

6

" "

July 27

-July I to Oct. I

Hyla cinerea

9

" "

July 2 to Oct. I

Bufo quercicus

17

" "

July 21

July 7 to Aug. 16

There are many discrepancies in this table. Of the first four species which
winter over we collected few transformed individuals. The earliest we reached
the swamp was April 24. All may begin to transform earlier than the table
shows. Doubtless the bulk of their transformation comes before the bulk of
transformation for the first five breeders of the spring, i. e. the February to
March 10 beginners. From Acris gryllus onward to the end of the table the
spawning order seems to have little relation to transformation dates. This is
more from insufficient data than anything else. In the table the least signifi-
cant item is the average date. In some cases it means little. From Acris
gryllus onward our transformation data come all from June i onward with one
exception and all average dates are in July whereas the first early breeders
have transformation from April i onward and averages in June or earlier.

If a composite curve were made of the 2 1 curves of transformation-sizes,
we would have a continuous record of sizes from 6 to 59 mm. The transforma-
tion sizes fall into four groups of sizes. (Table 1 8) The first group of 6-1 2 mm.
in size, (average 8-11, mode 7 to 11 mm.) comprises 6 species of four families:
The toads ( 5 w/omdae), namely oak toad and southern toad; two swamp cricket
frogs or chorus frogs {Pseudacris of Hylidae), little chorus frog, black cricket
frog; the narrow mouthed toad (Brevicipitidae), and the solitary spade-foot
iScaphiopodidae). The second group of seven species are from 9-23 mm. in
range, 12-20 mm. in average size, and 12 to 20 mm. in modal size. They are
all the remaining tree frogs ( Hylidae) . The third group of four species com-

Frogs of the Okefinokee Swamp

TRANSFORMATION SIZE

Table i8

Transformation size

Adult size

:, (mm.)

(mm.)

Smallest

Range

Averag

8 Mode

Largest
adult

breeding
adult

Bufo quercicus

6 - ID. 5

8

7

32

IQ

Bufo terrestris

6.5 10.5

8

8

88

42

Pseudacris ocularis

7 II

8

9

175

II-5

Pseudacris nigrita

(8? 12?)

30

21

Gastrophryne carolinensis

7 12

II

II

36

20

Scaphiopus holbrookii

8.5 12

10

ID

72

50

Hyla squirella

II 13

12

12

37

23

Hyla andersonii

II IS

12

12

47

30

Acris gryllus

9 IS

13

13

33

15

Hyla femoralis

10 15

13

13

40

23

Hyla cinerea

II-5 17

15

16

63

37

Hyla versicolor

13 20

16

16

60

32

Hyla gratiosa

18 23

20

20

68

49

Rana sphenocephala

18 33

23

25

82

49

Rana clamitans

20.5 33-5

25

24

75

52

Rana virgatipes

23 32

27

66

41

Rana aesopus

27 35

108

68

Rana septentrionalis

29 40

34

36

72

48

Rana heckscheri

31 49

35

33

125

82

Rana grylio

32 49

44

43

161

82

Rana catesbeiana

43 59

53

54

200

85

prise the southern meadow frog, green frog, carpenter frog and gopher frog.
They range from 18 to 35 mm., average 23 to 27 mm., mode 2 4 to 25 mm. Two
of these are transformers which winter over as tadpoles. The last group of
four frogs (Ranidae) comprise the winter tadpole group, 29 to 59 mm, in
range at transformation, average 34 to 53 mm., mode 33 to 54 mm.

The first 14 species are of species which normally transform the summer
the eggs are laid. Of the last 7 species (larger transformation sizes) six are of
the group which winters over as a tadpole.

Viewed from the point of transformation size and smallest breeding adult
in each species the interval order between ranges of transformation size and
smallest breeding size is as follows: (Table 19)

Table

19

Interval (mm.)

Interval (mm.)

Acris gryllus

0

- 6

Rana sphenocephala

16 -31

Pseudacris ocularis

5-45

Rana clamitans

18.5-31-5

Bufo quercicus

8

-II

Hyla cinerea

20 -25.5

Gastrophryne carolinensis 8

-13

Hyla gratiosa

26 -31

Hyla femoralis

8

-13

Rana catesbeiana

26 -42

Pseudacris nigrita

9

-13

Rana terrestris

3I-5-35-5

Rana septentrionalis

8

-19..

Rana aesopus

33 -41

Rana virgatipes

Q

-18

Rana grylio

33 -50

Hyla squirella

10

-12

Rana heckscheri

33 -51

Hyla versicolor

12

-19

Scaphiopus holbrookii

38 -41-5

Hyla andersonii

15

-19

General Discussion

59

Or if each species be considered from the point of view of least and greatest
transformation sizes, least breeding size and greatest breeding size, how many
times bigger will a little transformed anuran of each species have to grow to
attain least breeding size and greatest breeding size: (Table 20)

Tab

LE 20

To attain least

To attain greatest

breeding size

breeding size

Bufo terrestris

6.4-4 times

Bufo terrestris

13.5-8.4 tin

.Scaphiopus

Scaphiopus

holbrookii

6.0 -4. 18 times holbrookii

8.5 -6.0

Hyla cinerea

3.2 -2.2

Hyla cinerea

5

5 -3.6

Bufo quercicus

3.16-1.8

Bufo quercicus

5

3 -3

Gastrophryne

Gastrophryne

carolinensis

2.85-1.65

carolinensis

5

14-3

Hyla gratiosa

2.7 -2.12

Rana grylio

5

0 -3-3

Hyla andersonii

2.7 -2.0

Rana catesbeiana

4

6 -3-3

Rana sphenocephala

2.7 -1-5

Rana sphenocephala

4

6 -2.5

Pseudacris nigrita

2.62-1.8

Hyla versicolor

4

5-30

Rana heckscheri

2.6 -1.6

Hyla andersonii

4

3-31

Rana clamitans

2.6 -1.5

Rana aesopus

4

0-3.0

Rana grylio

2.55-1-6

Rana heckscheri

4

0-2.6

Rana aesopus

2.5 -1.9

Hyla femorahs

4

0-2.6

Hyla versicolor

2.5 -1.6

Hyla gratiosa

3

7-30

Hj^la femoralis

2-3 -1-53

Pseudacris nigrita

3

75-2.5

Hyla squirella

2.1 -1.77

Rana clamitans

3

65-2 . 2

Rana catesbeiana

2.0 -1,45

Acris gryllus

3

6 -2.2

Rana virgatipes

1.8 -1.3

Hyla squirella

3

3 -2.8

Rana septentrionalis

I. 75-1. 2

Rana virgatipes

2

9-2.0

Pseudacris ocularis

I .64-1 .04

Rana septentrionalis

2

5 -1.8

Acris gryllus

1.6 -I .0

Pseudacris ocularis

2

5 -1.6

RATE OF GROWTH

(PI. XVII)

Knowing the transformation size and average, mode and maximum adult
size we have certain starting points. But our conclusions on the number of
years of growth are based on presumptive evidence, namely on the meas-
urements of frogs taken at the same time or place, e.g. if at one time and place
we take a 14 mm. transformed frog, a 22 mm. frog, a 31 mm. frog, and a 40
mm. individual of Hyla femoralis, we might infer we had four age groups:
14 at transformation, 22, one year old, 31, two year old, and 40, three year old.
But what the resident naturalists can do is to mark frogs or keep them in
some pond inclosure and get the real growth of an individual or individuals.
My notes are to stimulate work along this line, little having been done in
this country.

Of course great variations might happened with some northern species
which might have extended periods of breeding, such as Bufo americanus,
Scaphiopus holbrookii or Rana clamitans, but in the south or southwest it
becomes more complicated with frogs which may breed from February or

6o

Frogs of the Okefinokee Sivamp

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General Discnssio7i 6i

March to November i or later. Some have assumed that if at outcoming in
the spring the collector find several separate size groups around certain means
we might have year groups. If however a species breeds from February to
December i he might have a variation from 4 months to 14 or 15 months.
One needs to know transformation sizes (range, average, mode), transforma-
tion periods (average, mode, range), breeding periods and countless other
factors to make a decent approach to a presumptive determination of year
groups. The following table is therefore provisional, based on what evidence
we have, and may be easily assailed or in part controverted but may lead to
long continued careful studies by subsequent workers.

In view of the fact that each of these series of presumptive ages was de-
termined from plotting the sizes of each species with no thought of previous
plottings for kindred species there are quite close parallels in this table. Still
one might ask how Rana sphenocephala might reach 78 mm. at the end of the
third year when R. clamitans takes 4 years according to the table? The
plotting for one or the other may be wrong or each individualistic. Rana
clamitans in the south is small for its species. The south is the optimum for
Rana sphenocephala.

In the same way Viosca has called my attention to the rapid growth of
bullfrogs in the south as contrasted to the north. The growth given for Ra7ia
catesbeiana is for the north, may be very presumptive and far from the truth
for Texan, Louisianan, Georgian and North Carolinian frogs. Bullfrogs I
have found breeding in February at San Antonio, Texas, but in Ithaca, N. Y.
never until June or July. The Rana catesbeiana figures in some ways do not
strictly belong in this table. (Table 21)

In Table 2 1 we have underscored the year each is held to breed. From this
we discover that Pseudacris ocularis, Pseudacris nigrita probably, and AcjHs
grylhis breed at i year old. These are the small tree frogs. Then comes the
oak toad (Bufo quercicus) and narrow-mouthed toads {Gastrophryne carolin-
ensis) breeding at 2 years of age. In the tree frogs four species {Hyla femor-
alis, Hyla squirella, Hyla andersonii, Hyla versicolor) apparently fall in the
two year old class, and three frogs (Rana virgatipes, R. sphenocephala and
Rana septentrionalis) also breed two years after transformation. This gives
representatives of four families, nine species in all, or about one-half of the
table maturing in 2 years or thereabouts. Two tree frogs ( Hyla cinerea and
Hyla gratiosa) breed as 3 year olds, and five frogs {Rana clamitans, Rana
aesopus, Rana heckscheri, Rana grylio, and Rana catesbeiana) fall in the same
class. The last group is one of the most precocious groups of early transfor-
mation, namely the southern toad and solitary spade-foot. They transform
quickly at small sizes, but have to grow apparently for 4 years before breeding.

In interpretation of age groups more chance for personal equation comes
in the early sizes. For example from transformation onward some would
make the frogs grow much faster the first year or so. Some no doubt would
merge my first year and second year group into one (namely yearhngs).
Herein is one of the crucial periods needing more study.

62 Frogs of the Okefinokee Swamp

FOOD

No phase might prove more profitable than the study of the food of these
species. Former associates of the author such as Drs. P. A. Munz and S. W.
Frost have pointed out how engaging this topic might be and how valuable
anura are in collecting the lowly and rare insects the entomologist might miss.
In this paper this topic is one of its weakest phases treated. Only three of the
Okefinokee forms have been carefully studied with Okefinokee material.
Dr. Vernon Haber published his interesting paper on "The Food of the
Carohna Treefrog, Hyla cinerea Schneider" (Journ. Comp. Psych. Vol. VI,
No. 2, Apr. 1926, pp. 189-220). Out of this study come G. Steiner's signifi-
cant "Some Nemas from the Alimentary Tract of the Carolina Tree Frog
(Hyla carolinensis Pennant) with Discussion of Some General Problems of
Nematology." (Journ. Parasitology, Sept. 1924, Vol. XI, pp. 1-32).

One never knows whither these food studies of frogs as well as other
groups will lead. In a somewhat similar fashion we have a by-product in the
study of the food of the oak toad (Bufo quercicus). Dr. Remington Kellogg
examined all our Bufo quercicus and Bufo terrestris from the Okefinokee
swamp and these studies are incorporated in his two forth-coming papers.
But some of the spiders of the Bufo quercicus food contents were submitted
to Prof. C. R. Crosby. He picked up one vial of spiders collected by Bufo
quercicus (Dr. T. Barbour collection) in Florida. He secured a new genus
and new species. The next vial were spiders collected by Bufo quercicus
(A. H. Wright collection) from Okefinokee swamp and the new genus from
Florida appeared and another new species of it. Professor Crosby gives it
the amusing title "A New Genus and Two Species of Spiders Collected by
Bufo quercicus (Holbrook)," but the title nevertheless expresses the real
situation. Other species ought to prove equally attractive forms for study.

ENEMIES

No article is more sought for or more relished as a food by a diversity of
animals from fish to man than frogs. The latter's defense consists in conceal-
ment and in the possession of poison glands in the skin, neither of which means
is aggressive in its nature. Insects and plants may prove a more constant
fare, but to any fair-sized animal a frog diet is one of the preferred menus, if
obtainable. Man is not content with cleaning up and draining the "frog holes"
or swampy stretches, but he kills the frogs at all seasons. He takes the frogs
when they congregate for breeding purposes and such a toll hardly accords
with the ideas of conservation held at the present day for birds or mammals.
The day may come when every farm will have its pond for cattle and hence
frogs, but swine are amongst their worst enemies. (For longer discussion of
this topic see the topic enemies under each species account.)

AFFINITIES

We are starting with the better known comparative characters of adults
and are adding a supplement out of our actual experiences with these frogs

Plate XV
Transformation

Transformed frogs. Lateral aspect.

1. Hyla andersonii, Lakehurst, N. J. July 5, 1923. x 0.75.

2. Hyla cinerea, Billy Id., Ga. July 25, 192 1. x i.o.

3. Hyla femoralis, Chesser Id., Ga. July 2, 1922. x i.e.

4. Hyla gratiosa, Chesser Id., Ga. July 27, 1922. x i.o.

5. Hyla squirella, Folkston, Ga. June 27, 1922. x 1.0.

6. Hyla versicolor, Chesser Id., Ga. June 29, 1922. x 1.0.

7. Acris gryllus, Chesser Id., Ga. July 8, 1922. x 1.0.

8. Pseudacris ocularis, Saddlebag Pond, Trail Ridge, Folkston, Ga. June 30,

1922. X I.e.

9. Scaphiopus holbrookii, two transforming frogs, Chesser Id., Ga. June 29, 1922

X 0.9.

10. Gastrophryne carolinensis, Chesser Id., Ga. June 25, 1922. x 1.0.

11. Bufo terrestris, Trader's Hill, Ga. June 24, 1922. x 1.0.

12. Bufo quercicus, Chesser Id., Ga. July 13, 1922. x 1.0.

Typical series, tadpole to transformed frog.

13-16. Hyla gratiosa, Chesser Id., Ga. July 25, 1922. Dorsal aspect, x 1.2.

lO

12.

14 15

Plate XVI
Transformed Ranas

1. Rana virgatipen, Lakehurst, N. J. May 26, 1924. \'entral aspect, x 1.3.

2. Raiia aesopus, Atlanta, Ga. August 27, 1922. Dorsal aspect, x i.o.

3. Rana grylio, Billy Id., Ga. April 28, 1921. Photographed with 4 for com-

parison. Lateral aspect, x i.o.

4. Rana clamitans, Billy Id., Ga. April 28, 1921. Lateral aspect, x 1.0.

5. Rana clamitans, South of Spanish Creek, Folkston, Ga. July 29, 1922. Lateral

aspect. X I.e.

6. Rana grylio, Billy Id., Ga. April 26, 1921. Ventral aspect x 1.2.

7. Ratia sphenocephala, Chesser Id., Ga. August 11, 1922. Lateral aspect, x 1.0.

8. Rana sphenocephala, Chesser Id., Ga. x 1.0.

9. Rana septentrionalis, Onekio, X. Y. July 5, 1923. Small specimen. Ventral

aspect, x I.e.
10. Rana septentrionalis. Onekio, X. Y. July 5, 1923. Large specimen. Lateral
aspect. X I.o.

Plate XVII
Chart of growth
Ranges of transformation sizes in comparison with the greatest mature sizes of
males and females of twenty species.

(measurements in millimeters)

O 10 ZO -JO ^ 50 fcO 70 60 go

IK IZO (30 l«0 150 IfcC 170

Bufo ,^«oc..
Bu[c lt,r«t,.s
Acr,.^,^llus.
Pieuddcn* n.g..t)
Reuaacr.iotuU'
HyU dnJc'tox.
h.,U c'.ne.pa

■,1a ^TiTiosi

yU scju.rella

Hyb vprs.colcv

nana sepTentrt-
ephenoceph-
Rana v.r^dt.pc^

General Discussion 63

in camp, in the field and with them ahve. One needs to gather much material
alive in its own haunt and seek it oneself over a period of time, to appraise
affinities and then he may be far amiss. Some of the evidences may be as
follows :

1. Adult characters. Some may think or hold too little attention has
been given comparative morphology and structural characters of the adults.
Heretofore these have been largely used and are the common presumptive
evidence already known and most easily sought with preserved material.
Had Holbrook known that A oris gryllus and Hyla crucifer laid single eggs
he would have been more confirmed in making them Hylodes gryllus and
Hylodes crucifer which he did on adult characters. When one knows Ra7ia
virgatipes lays different egg masses from Rana grylio then do the adult re-
semblances become so overpowering in assignment of relationship? If we
rehed on egg mass, R. virgatipes might be put far removed from R. grylio, yet
somewhere near R. grylio it doubtless belongs.

2. Voice characters of males. On the basis of sacs Bufo quercicus belongs
in a different group from Bufo terrestris, B. americanus, or Bufo fowleri, and
other characters bear out this judgment. Vocal sacs, however in the Hylidae
of Okefinokee are not so useful in separation of racial groups. In the Ranidae
the order of vocal sac development seems least in R. clamitans, R. catesheiana,
R. heckscheri, R. grylio, R. septentrionalis, and R. palustris. There is a
group with prominent lateral sacs, to wit, Rana sphenocephala, R. pipiens,
R. virgatipes, R. areolata, and R. aesopus. The first assemblage seems natural,
but does R. virgatipes belong in the second group on other grounds? The
second assemblage is as natural except for R. virgatipes. From another
vocal point of view, R. virgatipes and R. septentrionalis must be close because
each independently has been dubbed a Carpenter Frog.

3. Breeding maturity. On this basis we would put the small Bufo querci-
cus apart from Bufo terrestris. In the Hylidae, Acris gryllus, the species of
Pseudacris, Hyla squirella and Hyla femoralis would form a small sized
group; Hyla andersonii, Hyla versicolor, Hyla cinerea, and intermediate
group; and Hyla gratiosa, a large sized group. Or Rana virgatipes, R. sep-
tentrionalis, R. sphenocephala, Rana pipiens, and R. clamitans might make one
group and Rana areolata, R. aesopus, R. heckscheri, R. catesheiana, and R.
grylio another group. Are these groups good criteria of relationship in each
case? I fear not.

4. Breeding season. In the Ranas it would seem as if Rana sphenocephala
(also R. pipiens) and Rana aesopus (also R. areolata) fall in the very early
group, and Rana virgatipes, R. heckscheri, R. grylio, R. catesheiana, R. clami-
tans and R. septentrionalis form a later group. They seem natural groups,
but does R. palustris fall intermediate?

In the Hylidae, Pseudacris nigrita comes very early. Hyla gratiosa and
Hyla squirella and Acris gryllus, single egg layers, fall into early spring
breeders. Are the two Hylas related? The next group seems Hyla cinerea,
Hyla femoralis, Pseudacris ocularis, Hyla versicolor, Hyla andersonii, the
first and last possibly related, the second and fourth, but the third apart.

64 Frogs of the Okefinokee Swamp

o. Egg mass. In the frogs, Rana catesheiana, Rana grylio, (R. heckscheri),
R. clamitans lay films; Rana pahistris, Rana pipiens, Rana sphenocephala,
Rana aesopus, Rana areolata, Rana virgatipes and Rana septentrionalis lay
plinths or masses beneath surface of water. In the Hylidae, Hyla squirella,
Hyla gratiosa, Pseudacris ocularis, Hyla crucifer, Acris gryllus lay single eggs,
Hyla andersonii and Hyla cinerea are in an intermediate group, and Hyla
versicolor and Hyla femoralis lay surface films. Again we have recurring
some of the groupings of the other headings. Biifo quercicus and Bufo ter-
restris are separate groups entirely on this basis.

6. Individual eggs. In the Hylidae possibly Acris gryllus, Hyla crucifer
and Pseudacris ocularis should go together into one group of single eggs,
Hyla squirella and Hyla gratiosa into another, and Hyla cinerea and Hyla
andersonii into an intermediate group between single eggs and surface films.
Of the latter type we have Hyla femoralis and Hyla versicolor. In the Bufos,
Bufo quercicus belongs nearer Bufo compactilis or others of the southwest
than with B. terrestris which in turn is as near B. fowleri as B. americanus.
In the Ranas, Rana pipiens, R. sphenocephala, Rana aesopus, R. areolata, and
R. palustris fall together on this character, while Rana catesheiana and Rana
virgatipes have no inner envelope and R. grylio, R. septentrionalis and R.
clamitans have an inner envelope of jelly.

7. Larval period. The over-winter group of R. virgatipes, R. clamitans,
R. septentrionalis, R. heckscheri, R. grylio, and R. catesheiana, seems a natural
group. They are later breeders than the short larval period group of Rana
pipiens, R. sphenocephala, R. aesopus, R. areolata, and R. palustris which
breed earlier except R. palustris. The Hylids, nine species, separate in no
such clearly cut fashion.

8. Mature larvae. The tadpoles of R. pipiens, R. sphenocephala, R.
aesopus and R. areolata fall more or less together as do those of the wintering
tadpole group of six species {R. clamitans, R. catesheiana, R. grylio, R. heck-
scheri, R. virgatipes, and R. septentrionalis.) In the Hylids, Acris appears as
much a Ranid as a Hylid in character. Hyla crucifer and the species of
Pseudacris apparently fall together into another group. Hyla gratiosa, Hyla
andersonii, and Hyla cinerea seemingly are another group. And Hyla
femoralis, Hyla versicolor and Hyla squirella form a fourth group.

9. Transformation period. The six Ranas {R. clamitans, R. grylio, R.
heckscheri, R. virgatipes, R. catesheiana, R. septentrionalis) begin to transform
early the following season. Later come Rana pipiens, R. sphenocephala or
Rana aesopus and Rana areolata. In the Hylids Pseudacris nigrita transforms
early, the others much later and over a larger period. So in the Bufos, Bufo
terrestris transforms early and Bufo quercicus much later.

10. Transformation size. Apparently this gives little clue to relationship.
The group of Rana sphenocephala, Rana pipiens, Rana aesopus, Rana areolata
may have smallest transformers (the first two) or larger transformers (the
last two) or in the same way these groups may attain medium size or large
size as adults. In the same way Rana virgatipes is small in transformation
and adult size in its group {R. grylio group) which in the main is large in
these particulars.

General Discussion 65

11. Transformation structures. Often at transformation it is extremely
difficult to determine the Hylids or tree frogs. They are frequently each of
them green at this stage. The taking on of their color pattern from trans-
formation onward may often give a clue to relationship. For example, on this
basis one might be tempted to place Hyla cinerea, Hyla andersonii, and
Hyla gratiosa together.

HYPOTHESES

This topic is dangerous. It might give a fine chance for pleasurable
writing. One might develop it with increasing confidence as one grew older.
None of the following suggestions do we urge and their tentative characters
merely show some habits of mind which have pervaded previous attempts
at the relationships, distribution and origin of American biota. I hope no one
will expend most of their energy on this part of our paper and tilt any hypothe-
sis hard until he or she has labored industriously on the forms in question.

Some of these fancies are: i. Is it worthy of consideration that those
forms of Europe which Boulenger places below 4,200 feet elevation (namely,
Rana agilis, Rana esculenta, Bufo calamita, Hyla arborea, Pelobates fuscus,
and possibly others) might have counterparts in our Atlantic coast region
in Rana sylvatica, Rana virgatipes, Bufo fowleri, Hyla andersonii, Scaphiopus
holbrookii, etc.? Three of them are yet unrecorded from Okefinokee region
and one not common and one on the outside of the swamp.

2. Is the Okefinokee region near the center of distribution of the forms
of the United States of America with its eight Hylids (2 Pseudacris, i Acris,
5 Hyla), and three more possible Hylids, (2 Pseudacris and Hyla andersonii)
or six Rana species or two diverse Bufos or i Scaphiopus and i Gastrophryne —
18 species with 4 or 5 more possible, or 22 or 23 in all? For example, would it
be safe to say the region of the south eastern United States is the center of
the single laying Hylids, namely Hyla squirella, Hyla gratiosa, Acris gryllus,
Pseudacris ocularis, and that from one of these came the lone northern species,
Hyla crucifer? Or from a Rana grylio-\ike form came the more widespread
northern R. catesbeiana, R. virgatipes, and Rana septentrionalis, all very
aquatic? Or from Rana sphenocephala came the widespread northern Rana
pipiens, or from Ra7ia aesopus, the more widespread northern Rana areolata?

3. Or are some of the Okefinokee species so hmited in distribution
(South Carolina to Louisiana, or less) to be merely Sabalian representatives
of northern species? Shall we call the aberrant Pseudacris ocularis a Sabalian
offshoot from the northern Hyla crucifer, Rana grylio a bigger Rana virga-
tipes, Rana heckscheri a bigger Rana clamitans, Hyla gratiosa a bigger Hyla
ciner'ea?

4. Did most of our southeastern forms come from the southwestern or
Mexican center, another favorite method of explanation? Did Bufo quercicus
come from Bufo compactilis, Bufo cognatus or some southwestern form, or
Bufo terrestris from Bufo valliceps'^ Is Hyla squirella of the Hyla eximia,
Hyla regilla group? Or are Hyla versicolor, Hyla avivoca, Hyla phaeocrypta,
Hyla femoralis from a Hyla arenicolor center? Is it reasonable to hold that

LU LIBRARY

66 Frogs of the Okefinokee Swamp

the center of Scaphiopus (S. hammondi, S. hurteri, S. couchii) is southwestern
and that *S. holbrookii came from *S. couchii? Or are the two diverse Pseudacris.
lines: P. nigrita (clarkii) and P. ornata the center of all the others unless
it be P. ocularis? Or is Gastrophryne carolinensis the one eastern fringe
species from a Mexican center with several species?

5. Or must we think of some of the southwestern forms as outliers from
an eastern center? Hyla arenicolor a sole representative of Hyla femoralis
and Hyla versicolor? Or is Hyla regilla the one western approach to Pseu-
dacris species?

6. Are Hyla gratiosa, Pseudacris ocularis, Bufo quercicus, Rana heck-
scheri, archaic or primitive relicts of an old faunal center in central Florida
or elsewhere in the southeast comparable to the peninsular forms such as
Neoseps, Rhineura and several snakes?

7. Or have some of these had ingress quite recently or accidentally,
possibly like E. ricordii, geckos, etc.?

8. Are no certain Nearctic or Holarctic types like Rana sylvatica, or
some Pacific Ranas or Ascaphus truei represented in the Okefinokee region?
Just as in mammals, birds, etc. slight if any true upper Austral or Transition
Zone forms appear.

9. Are the Atlantic coast forms like Bufo fowleri, Hyla andersonii, Rana
virgatipes absent or scarce just as Chrysemys picta, Clemmys muhlenhergii,
etc. are absent, Clemmys guttata very rare, etc., i.e. is the Atlantic coastal
influence attenuated almost to nothing and Sabalian conditions predominant?

10. These queries might be expanded with no positive statements made
but enough herein appear to show the bases for these imaginative indul-
gences or speculations.

SUMMARY OF AFFINITIES
In a sentence we might venture the following tentative assertions regarding
each species of this study.

Scaphiopus holbrookii falls in relationship with Scaphiopus couchii, S.
hurteri and less so with S. hammondii.

Bufo quercicus belongs near some western or southwestern forms such
as Bufo compactilis and others.

Bufo terrestris may be of the Bufo fowleri, B. americanus group or
related to Biifo valliceps.

Acris gryllus, the most aberrant Hylid, possibly more Ranid (see
hind femur like R. grylio or R. virgatipes or tadpole) but possibly near
Hyla crucifer or Pseudacris ocularis.

Pseudacris nigrita, much Uke P. triseriata, P. feriarum, etc., and pos-
sibly of a different line from Pseudacris ornata.

Pseudacris ocularis may be as related to Hyla crucifer or Acris gryllus
as any of the Pseudacris nigrita assemblage.

Hyla andersonii probably falls close to Hyla cinerea group or possibly
near Hyla squirella.

General Discussion 67

Hyla cinerea may be the same as Hyla c. evittaia and close to Hyla
andersonii or possibly to Hyla gratiosa.

Hylafemoralis is the Hyla versicolor representative of the piney woods.

Hyla gratiosa may be apart but possibly may fall with Hyla cinerea
or Hyla andersonii.

Hyla squirella. Does it fall with Hyla crucifer and .4cm gryllus, or
Hyla cinerea {cinerea, andersonii, gratiosa) group, or Hyla versicolor
( H. arenicolor, femoralis) group?

Hyla versicolor has as its nearest relative in Okefinokee, the piney
woods treefrog Hyla femoralis.

Ratia aesopus is close to Rana areolata. These in turn are close to
R. sphenocephala and R. pipiens.

Rana catesbeiana doubtless has its nearest relative in R. grylio or
R. clamitans.

Rana clamitans is probably nearer Rana heckscheri than R. catesbeiana
or Rana grylio.

Rana grylio is a bullfrog, doubtless belonging in the R. catesbeiana
group, yet it has affinities with R. clamitans or possibly R. virgatipes.

Rana heckscheri doubtless comes closest to R. clamitans though more
must be known to place it carefully.

Ra7ia septentrionalis is a puzzle. Is it close to R. virgatipes or near
R. clamitans, or a holoarctic form from Eurasian stock like some north-
western species?

Rana sphenocephala is a southern Rana pipiens.

Rana virgatipes may be a relative of R. grylio or close to R. septen-
trionalis.

Gastrophryne carolinensis in tadpoles, eggs, etc. is different from G.
texensis. Possibly this or a Mexican species may be its nearest form.

SUGGESTIONS FOR LOCAL OR RESIDENT NATURALISTS AND
COLLECTORS IN THE SOUTHEASTERN UNITED STATES

General:
Topics in these life histories which in general are incomplete, fragmentary^
or weak are:

1. First appearance. 9. Growth studies on marked indi-

2. Migrations to breeding places. viduals, not presumptive.

3. Range of cross-mating. 10. Food.

4. Egg-laying process. 11. Autumnal disappearance.

5. Range in egg complements. 12. Hibernation.

6. Hatching period in the field. 13. Sexual discrimination in mating as

7 . Variations in mouthparts of revealed in cross, abnormal, uni-
tadpoles sexual embraces.

8. Larval periods made more exact.

68 Frogs of the Okefinokee Swamp

Special:
Scaphiopus holbrookii

1. Experimental zoologists should canvass why its tadpoles develop into
frogs in 17-30 days.

2. Why are sizes from 12-51 mm. scarce in collections?

3. Even presumptive growth data is scanty.

4. Food might prove very suggestive. Entomologists collect more fre-
quently early night flying insects.

5. Post breeding studies to hibernation are fragmentary. This is the
age-old question about spadefoots.

6. Why do S. holbrookii and S. couchii, large adults have small tadpoles,
and transform at small sizes, and S, hammondii a small ad^lt, trans-
form at a large size? •

Bufo quercicus

1. More observations on egg-laying process are neded.

2. The mature tadpole needs to be better described.

3. Transformation data scanty.

4. Bufo quercicus as an insect collector of small forms (see Dr. Remington
Kellogg's and Professor C. R. Crosby's studies of this form).

Bufo terrestris

1. Tadpoles of Bufo terrestris need to be carefully described.

2. Are its affinities with B.fowleri and B. americanus or with B. valliceps,
etc.?

3. Discrimination or non-discrimination of males at the breeding season.
Acris gryllus

1. How distinct are Acris crepitans and Acris gryllus?

2. Do 15 mm. transformed frogs breed at once or are they from 9 mm,
transformed sizes?

3. Why superlarvation in Acris? Does it imply diverse larval periods?

4. Does it regularly lay single eggs or masses?
Pseudacris nigrita

1. We missed its life history completely. Every topic in our outline for
each species needs attention.

2. Is it really different from P. triseriata, P. feriarum, P. septentrionalis
and (old) P. clarkii?

Pseudacris ocularis

1 . Does it belong in Pseudacris?

2. More evidence on breeding season.

3. Egg laying in field.

4. Larval period problematic.

5. Probable differences between tadpoles of P. ocularis, P. nigrita, and
P. ornata not known. Lends an uncertain element in identification of
Pseudacris larvae of southeastern U. S.

6. This mite of a frog ought in its food to collect unusual forms for
entomologists and parasitologists.

General Discussion 69

Pseudacris occidentalis
Pseudacris ornata.

1. One species or two?

2. In either case complete life histories needed.

3. Will the toadlike P. ornata be as diverse as P. ocularis or P. triser-
iata?

Hyla andersonii

1. A better color description from life of the female is desirable.

2. The breeding sizes (30 mm.-4i mm. males and 38-48 mm. females)
are apparently lacking in breeding females below 38 mm. and breeding
males below 30 mm.

3 . Is Hyla andersonii in other parts of its range always a sphagnaceous
breeding frog? (See R. virgatipes)

4. What is the hatching period in life?

5. Larval period needs more evidence.

6. Transformation period and size needs more observation.

7. Even presumptive growth impossible because there are so few speci-
mens in collections between 15 mm. and 35 mm.

Hyla cinerea

1. Is it primarily a surface-film or submerged-strewn egg breeder?

2. After the discovery (1922) of R. heckscheri young tadpoles, greater
refinement of H. cinerea description is needed. Few H. cinerea tad-
poles found in 1922.

3. More transformations are needed.

4. Rate of growth problematic in intermediate sizes until more material
appears.

5. Study transformed green tree frogs of H. andersonii, H. cinerea, H. c.
eviiiata, H. squirella, and H. gratiosa for clues to relationships when
they begin to assume their adult patterns.

Hyla femoralis

1. This common Sabalian form could be used for cross breeding experi-
ments and possibly hybridization with other Hylas.

2. The larval period needs to be better determined.

3. This denizen of the high long leaved pines would be a good subject for
food studies. It would yield interesting material of the upper levels
of the pine barrens.

4. Abnormal variations in its tadpoles might be interesting to follow.
Hyla gratiosa

1 . What is the post and pre-breeding habits of this species?

2. Why two dissimilar calls in this species?

3. Egg-laying process in laboratory and in field needs to be watched.

4. Intermediate sizes from 25-50 mm. are badly needed in collections for
growth studies.

Hyla squirella

I . Why is this Hyla more of a house treef rog or found more about human
quarters, habitations, barns, cultivated yards, etc.?

-JO Frogs of the Okefinokee Swamp

2. Why do so many breeders choose such temporary pools?

3. Variations in the coloration are worthy of detailed study.

4. Does this species occasionally lay a surface film different from its
normal singly laid eggs?

5. Larval period needs more attention.
Hyla versicolor

1. Why is this species not within the swamp?

2. Are these more marked in color than the northern forms?

3. Are the breeding periods of the north and south synchronous?

4. Why did the species average smaller in adults, tadpoles and trans-
formation in southeastern United States?

Rana aesopus

1. Are R. capito and R. aesopus one?

2. Mating of the species unobserved.

3. Transformation notes in general are much to be desired.

4. Intermediate sizes from 35-48 mm. are very scarce.

5. What do the younger frogs do?
Rana clamitans

1. Not studied in as great detail because of northern notes on the species.

2. Why are breeding frogs much smaller than in the north and trans-
formation at a smaller size?

3. Hatching period in field much desired.

4. Data on larval period needed.
Rana grylio

1. The egg laying has not been observed.

2. No field mated pairs have been taken.

3 . Hatching records are not numerous.

4. The larval period is barely outlined.

5. What are its relationships in range to that of R. catesbeianal
Rana heckscheri

1. Why has it been overlooked so long?

2. Intermediate sizes from transformation (31-49 mm.) to 82 mm. are
needed.

3. Mating amplexation and habits not observed.

4. Method of ovulation and eggs not described.

5. Is its larval period one or two years?

6. Are albino tadpoles not infrequent in this form?
Rana septentrionalis

1. The breeding period needs to be outHned in better fashion.

2. Actual mating and amplexation we have not observed.

3. What is the normal hatching period?

4. More details of the variations in tadpoles are needed.

5. More positive evidence on larval period should be secured.

6. How long after transformation does or might a frog keep its tail?

General Discussion

71

Rana sphenocephala

1. What are the positive distinctions between R. sphenocephala and
R. pipiens?

2. In the southeast does R. sphenocephala breed ahnost every month of
the year?

3. Differences in young stages in tadpoles of i?ana aesopus and Rana
sphenocephala.

4. Are albino eggs often laid in coffee colored water?

5. Hatching period and larval period may be different with eggs laid
almost every month of the year.

Rana virgatipes

1. Mating amplexation never observed.

2. Eggs needed to be identified other than b}^ ehmination. Check
need from a mated pair.

3. Why have R. virgatipes and R. septentrionalis submerged egg masses?

4. The tadpole description needs supplementation and the tadpole
period data is meager.

5. Transformation data is also fragmentary.
Gastrophryne carolinensis.

1. The vocal powers of the male and the process might be an interesting
study.

2. The egg laying process has not been observed.

3. The food of the queer tadpole ought to be interesting.

4. Value of smooth or rough skin in taxonomic analyses of narrow-
mouthed toads.

5. The spiracle, tadpole shape, intestine, absence of normal mouthparts,
etc. of tadpole might be made a special study.

Bufofoivleri, Hyla avivoca, Rana catesheiana

1. Hyla avivoca will doubtless be found eastward to Georgia and far-
ther. Its life history is unknown.

2. The southeastern distribution of Bujo fowler i in distinction from
Bufo americanus and Bufo terrestris and of Rana catesheiana in dis-
tinction from Rana grylio and Rana heckscheri is needed.

GENERAL ACCOUNTS OF SPECIES
Scaphiopus holbrookii (Harlan)

(PI. I, Fig. 4; IV, Fig. 2; VI, Fig. 2; X, Fig. 12; XIV, Fig. 8; XV, Fig. 9;
XVII; XVIII; XIX; Text Fig. i, 5)

COMMON NAMES

Spade-foot. Hermit Spade-foot. Hermit Spade-foot Toad. Spade-foot
Toad. Hermit Toad. Solitary Spade-foot. Solitary Toad. Holbrook's
Spade-foot. "Storm Toads." "Storm Frogs."

RANGE

Check list. "Type Locality South Carolina. Range: Eastern States,
Mass-achusetts to Florida, west to Louisiana, Texas to Arkansas." Stejneger
and Barbour's Check List 1923, p. 23.

Supplementary records. Dr. Holbrook (1842, Vol. IV, p. iii) writes
"Its range is more extended than I first apprehended. It is found in Carolina
and Georgia. Dr. Troost sent me a specimen from Tennessee, and Dr. DeKay
has observed it in the State of New York." In 1849 (Appendix, p. 15) Hol-
brook gives Scaphiopus holbrookii as a frog of Georgia. Dumeril and Bibron
(1841, p. 474) expect it to be discovered "sur d'autres points des Etat-Unis."

In 1842 DeKay (p. 66) writes "We have now the pleasure to include it in
the Fauna of New York. Specimens of this animal were found by Mr. Hill,
in a garden near Clarkstown, Rockland County. . . , Dr. Pickering, I
learn, has recently seen it in the neighborhood of Salem, where they appear
in great numbers, at distant periods, after rains of long continuance."

The same year, 1842, DeKay was publishing, Andrew Nichols (1852,
pp. 113-117) of Massachusetts was making the first extended descriptions of
their habits. "Holbrook, in his N. A. Herpetology, Vol. I, pp. 85-87, says that
he found these reptiles in three states only, viz : Georgia, South Carolina, and
Tennessee — that they go in the water only in the breeding season, that which
he observes in the spring; and that they live in holes in the ground of about
six inches in depth, excavated by themselves — never coming out of these,
except during the night and after heavy rains. This explains the mystery
of their sudden appearance and disappearance, as above mentioned. It would
also seem that they are Southern reptiles; — chilled by our northern climate,
they want a more genial climate to celebrate their nuptials ; and thus without
a suitable pool to receive their spawn, year after year in this instance trans-
pired, until a summer freshet filled their native habitat sufficiently."

"I have some reasons to conjecture that other colonies of these frogs exist
in New England. An intelligent farmer of Topsfield (Mass.), to whom I
showed my specimens, and related the foregoing story, told me he had several

73 •

74 Frogs of the Okefinokee Swamp

times heard a similar croaking in a temporary pond of water near his dwelHng,
but he never went to see from whence the noise proceeded. Something of the
same kind has been noticed in Framingham, in this State, by a student of
medicine, who relates that he and a friend of his were kept awake on the
identical night, June i6th, 1842, by the noise of frogs or something of the
kind in a ditch of water near his lodging; that they went out to discover
what they were, but on drawing near the place, the noise or cry, which re-
sembled that of young crows, suddenly ceased, and nothing was to be seen."

In 1868 C. C. Abbott in New Jersey found that "This little frog is not
abundant, and is generally met with when found, in the Southern counties
of the State."

In 1909 Julius Hurter and John K. Strecker, Jr., record it in Green way,
Arkansas.

These previous records are not exactly supplemental but intended to show
that quite early the extremes of its range were fairly well known.

In May, 1922, in Florida, "Richard Deckert (1922, p. 88) ran over one at
Homestead, in the evening on the return trip." In 1923 A. H. Wright and
A. A. Wright report this species from the Okefinokee region, Georgia and
Florida. The same year Director T. VanHyning (1923, p. 68) of the Florida
State Museum reports two found at night, March 18, at Gainesville, Florida.

In the summer of 1926 A. I. Ortenburger (1927, p. 46) in Western Okla-
homa reports "Scaphiopus holbrookii holhrookii (Harlan) (in) Cimarron County
5 miles north of Kenton." In 1926 Wright (p. 83) gives this species as a
Lower Austral species and gives its range as Massachusetts to Florida. The
reason for the present author limiting it to Florida and not extending it to
Arkansas and parts west of Florida is not now apparent.

Local Okefinokee records. In 192 1 we were in the center of the swamp and
did not record this species. Mr. Matt Hickox told us of a frog different from
any we had on Billy's Island. His description of a special frog or toad which
came at heavy rainy periods we suspected to be Scaphiopus holhrookii. He
said it occurred in the vicinity of Braganza on the northeast border of the
swamp.

In 192 1 Mr. Harper heard from Mr. Dave Mizzell (on Folkston-Moniac
road) who told him of a frog "called 'storm frog' because he appears when
there are storms or floods. It hollers wank! It folds up so that you can't
find its legs. It is larger than the 'hop toad'. One was dug up in a potato
field."

In 1922 immediately after our arrival on Chesser Island on June 14, Mr,
Ridley Chesser told us of frogs he heard near their barn that were neither
carpenter nor gopher frogs. We suspected it of being spade-foots and on
Chesser Island. A week later, June 22, in one of the deep pits at Camp
Pinckney, Folkston, Ga., we found little else than spade-foot tadpoles. The
pit was heavily shaded by oaks. The following day, June 23, we recorded
tadpoles in a pond the Trader's Hill side of Brook's R. R. one mile from
Trader's Hill. On June 27 we make the following note that "because of the
discovery of these tadpoles in two different places this species seems a com-

Scaphiopus holhrookii 75

monly distributed form in these parts." We found several people who knew
about ''storm toads" or "storm frogs."

Then on Aug. 16-17, 1922, near Hilliard, Florida, we camped beside a spade-
foot congress in open cut -over pine-barrens, flooded because of an immense
rain just preceding.

GENERAL APPEARANCE

This phase has aroused as much interest as any topic treated by authors-
Its discoverer. Dr. Holbrook, (1842, Vol. IV, pp. 109-111) records "This
singular animal approaches nearly to the toad in form." "This is a strange
animal — an odd mixture of toad and frog, having the teeth of one, and the
rudimental post-tympanal glands of the other ; it approaches, however, nearest
the toad in its form and habits, . . . ." "The skin is very delicate, and though
warty or granulated after exposure, when first taken from its hole the Scaphio-
pus presents the etiolated appearance of a real subterranean animal." "Char-
acters. Back olive coloured and somewhat warty, with two lines of pale yellow
extending from the orbits to the vent, beneath yellowish white."

Harlan (1835, pp. 105, 106) says "This new species, first figured and des-
scribed by Dr. Holbrook, (vide his valuable work on North American Rep-
tilia) possesses very peculiar characters, displaying, in its external configura-
tion, a strange mixture of the toad and frog. It has the contracted form of the
first, with small typmanum, and rudimentary super-tympanal warts, without
visible pores, and small warts disseminated over the back of the head ; possess-
ing palatine teeth and serrated maxillae, like the frogs, and like the Rana
cultripes of Cuv. is remarkable in .possessing a rudiment of the sixth finger,
covered by a sharp horny plate, . . . ."

DeKay (1842, p. 66) gives as its "Characteristics. Ash grey, with two
yellow curved lines from the eyes, dilated and subsequently united at the vent.
Length two inches."

C. S. Brimley (1907, p. 157) gives upper jaw with teeth. Paratoids
present. Hind feet webbed. Heel with a flat sharp edged spur" as enough to
distinguish the Solitary Spadefoot. He uses the same characters in his re-
vised key (1926, p. 80),

MEASUREMENTS
(Recent Material)
//ead to angle of mouth 1.33 (28 mm.)— 1.36 (44 mm. ? ) — 1.35 (56 mm.cj^) —
1.27 (56 mm. 9 ) — 1.27 (68mm.c/') — 1.33 (68 mm. 9 ) in width of head; head to
rear of tympanum 1.09 — i.ii — 1.19 — 1.09 — 1.07 — 1.12 in width of head ; head
to angle of mouth 3.1 — 2.99 — 3.02 — 3.1 — 3.09 — 3.24 in length of body; head to
rear of tympanum 2 .54 — 2 .6^2 .66 — 2.66 — 2.66 — 2.61 — 2 . 7 2 in length of body ;
snout — .54 — .58 — .71 — .60 — .66 — .70 in first finger; snout .54 — .62 — .57 — .56
— .58 — .64 in fourth finger; snout .36 — .29 — .47 — .40— .37 — .34 in first toe;
eye 1.1—1.41— 1.5— 1.33— 1.5— 1.43 in snout; eye .4— .66— .71— .73— .81—
.75 in tympanum; eye .6 — .825 — 1.06 — .80 — i.o — i.oinfirst finger; tympanum
5-5 — 4-0 — 4-2 — 3.4 — 3-4 — 4-0 in intertympanic width; tympanum 2.75 — 2.25
— 2.1 — 1.8 — 1.84 — 1.75 in snout; internasal width 1.16 — 1.25 — i.o — i.o — 1.8

76 Fi'ogs of the Okefinokee Swamp

— i.o in upper eyelid width; interorbital width .77 — .77 — .625 — .75 — .72 — .68
in upper eyelid width; interorbital width .66 — .61 — .625 — .75 — .66 — .68 in
internasal width; interorbital width 2.44^ — 2.44 — 2.6 — 2.37 — 2.5 — 3.0 in inter-
tympanic width.

Forelimb: Forelimb 1.86 — 2.0 — 2.43 — 2.0 — 1.74 — 1.7 in length of body;
forelimb 2.13 — 2.04 — 2.39 — 1.75 — 1.87 — 1.7 in hind limb; first finger 1.5 —
1.7 — 1-33 — 1-5— I-3I — 1-75 in third finger; second finger 1.5 — 1.7 — 1.33 — 1.63
— 1.4 — 1.75 in third finger; second finger i.o — i.o — i.o — 1.09 — 1.06 — i.o in
first finger; third finger .88 — .59— .75 — .72 — .95 — .64 in second toe; fourth
finger 1.5 — 1.88 — 1.66 — 1.63 — 1.5 — 2.0 in third finger; fourth finger .66 — .55 —
.825 — .72— .64 — .57 in first toe; internasal width 1.0 — 1.25 — 1.5 — 1.0 — 1.33 —
1.4 in first finger; internasal width 1.0 — 1.25 — 1.5 — .91 — 1.25 — 1.25 in second
finger; internasal width 1.5 — 2.12 — 2.0 — 1.5 — 1.75 — 2.5 in third finger; inter-
nasal width 1.0 — 1. 12 — 1.2 — .93 — 1. 16 — 1.27 in fourth finger.

Hindlimb: length 1.14 — 1.02 — .99 — .87 — 1.07 — 1.0 in hind limb; tiba —
2.54^ — 2.83 — 2.8 — 3.2 — 2.83 — 2.95 in length; tibia 2.9 — 2.9 — 2.75 — 2.8 — 3.04
— 2.95 in hind limb; tibia 1.36 — 1.42 — 1.15 — 1.6 — 1.62 — 1.74 in fore limb;
tibia 1.09 — 1.06 — I.I — 1.08 — 1. 16 — 1. 13 in hind foot; first toe 2.0 — 2.0 — 1.5 —
1.625 — 2.22 — 2.25 in second toe; first toe 3,0—3.6 — 2.4 — 2.75 — 3.33 — 3.25 in
third toe; first toe 4.5 — 5.0 — 3.3 — 3.5 — 4.9 — 4.75 in fourth toe; first toe 2.5 —
2.8 — 2.125 — 2.0 — 2.7 — 2.62 in fifth toe; second toe 1.5 — 1.8 — 1.6 — 1.7 — 1.5 —
1.44 in third toe; second toe 2.25 — 2.5 — 2.2 — 2.15 — 2.2 — 2.1 in fourth toe;
second toe 1.25^ — 1.4 — 1.33 — 1.3 — 1.25 — 1.16 in fifth toe; third toe 1.5 — 1.4 —
1.37 — 1.27 — 1.46 — 1.46 in fourth toe; third toe .825 — .77 — .83 — .77 — .8 — .80
in fifth toe; fourth toe 1.33 — 1.32 — 1.25 — 1.35 — 1.27 — 1.36 in hind foot with-
out tarsus; fourth toe 1.2 — 1.24 — 1.2^1.25 — 1.09 — 1.21 in tibia; fourth toe
1.66 — 1.76 — 1.4 — 2.0—1.68 — 2.1 in fore limb; fifth toe 1.8 — 1.8 — 1.65 — 1.64 —
1.76 — 1.8 in fourth toe; internasal width — .66 — .625 — 1.0 — .66 — .75 — .74 in
first toe; internasal width 1.33 — 1.25 — 1.08 — 1.5 — 1.66 — 1.4 in second toe;
internasal width 2.0 — 2.25 — 2.4 — 1.83 — 2.5 — 2.2 in third toe; internasal width
3.0 — 3.15 — 3.3 — 2.33 — 3.66 — 3.4 in fourth toe; internasal width 1.66 — 1.75 —
2.0 — 1. 41 — 2.08 — 1.9 in fifth toe.

HABITAT

Colonel Nicholas Pike (1886, p. 213) holds 'Tt must, of course, be hunted
for on secluded places and woody hillsides, but I will venture to say even the
most knowing, in nine cases out of ten, will only find a Spadefoot by accident."

FIRST APPEARANCE

Holbrook places this in early March. Many authors have them appearing
from April onwards.

On December 2, 1868 Dr. J. A. Allen (1868, pp. 186 & 187) speaks of their
appearance at both Springfield and Cambridge, Massachusetts. He says:
"On hearing the very pecuhar notes of this species six years since, at the well-
known locality near the Botanic Garden in Cambridge, I recognized it at once
as being something I had heard occasionally at Springfield. It was not, how-

Scaphiopus holhrookii 77

ever, till May 27th, 1866, that I happened to be fortunate enough to obtain
specimens! I found two in a path, after heavy rain, several hundred yards from
any permanent pool; and, during the day, several pairs spawning in different
small transient pools, though at this time they appeared sparingly. I also
heard them in Chickopee the following day, five or six miles distant from the
first locality.

"In 1863, after unusually heavy rains towards the close of June, they came
out in immense numbers, the transient pools formed by the heavy fall of rain
seeming to be full of them; but they were heard only from i P. M. till about
3 A. M., of the following night; and being confined to the house by illness I
failed at this time to obtain specimens. Not having been in Springfield at the
proper time since, these are the only instances of their occurrence known to
me. Specimens collected here in 1866 were deposited by the writer in the
Springfield Natural History Museum, and in the Museum of the Essex
Institute.

"The character of the season seems greatly to determine the time of the
appearance of these animals, for they rarely, if ever, appear except after long
continued rains, during which the fall of water has been sufficient to saturate
the ground thoroughly, and to form pools in situations ordinarily dry. If the
spring opens with heavy rains, it is not uncommon for them to appear during
the last of March, or early in April, often before the snow is gone ; if dry they
are not seen till later, and if no heavy rains occur during the spring or summer
months, as sometimes happens, they are not seen at all that year. If suffi-
ciently heavy rains occur in May, or even in June, or in July, and not pre-
viously, as happened one year at Springfield, they may be expected even then.
The present year, remarkable for its wetness, seems fully to demonstrate this,
no less than four sets of the Scaphiopus having been observed at Cambridge.
The first, few in numbers, appeared during the few warm days that occurred
about April ist; the second, much more numerous, April 15th; the third May
14th, and the fourth May 22d; each during, or immediately following, a very
heavy falls of rain. They were observed not only at the old locality near the
Botanic Garden, but in several others, including the pond west of the Museum
of Comparative Zoology, and another east of it, formed by the temporary in-
undation of the marsh in Mr. Norton's woods. It takes but a few weeks for
the young tadpoles to mature, the eggs generally hatching in from five to seven
days, and the young being ready to leave the water in about three weeks;
yet the pools selected by the Spadefoot for the home of their off-spring often
become dry before the tadpoles are fully grown, and they consequently
perish, — so that ordinarily but a small part mature. The wetness of the
present year was not only favorable for their spawning, but also for the devel-
opment of the young, so that the increase of the Spadefoots must have been
unusually large."

GENERAL HABITS

Holbrook (1842, p. iii) holds that "it never ventures in water except at
the breeding season; it lives in small holes about six inches deep, excavated by
itself in the earth, which for a long time I mistook for holes of insects; here it

78 Frogs of the Okefinokee Swamp

resides, like the ant-lion, seizing upon such unwary insects as may enter its
dwelling. It never leaves its hole, except in the evening, or after long con-
tinued rains. It shows great dexterity in making its dwelling, sometimes us-
ing the nates, and fastening itself by the spade-like process; at others it uses
the legs with these processes, like a shovel, and will in this way conceal itself
with great rapidity. In progression its motions are not very lively, and its
powers of leaping but feebly developed."

Harlan (1835, p. 106) thinks the spade "must materially aid the animal in
climbing steep and slippery ascents. Habits are said to be peculiar; about the
size of the Esculent frog. Inhabits South Carohna."

DeKay (1842, pp. 66, 67) holds "It lives in small holes, in damp earth, a
few inches below the surface, which it excavates with great ease by means of
its spadehke processes. In these holes it hes in wait for such insects as may
approach, and I suspect can spring forth to seize whatever may be passing in-
cautiously near its hiding place. I remarked, at least in those which I had
alive, that it leaped with great apparent ease to a considerable distance. To
judge from those in my possession, although completely identical with the
solitarius, I should be disposed to believe that our northern variety is less
brilliant in its markings, and its general color is of a more grave and sombre
hue."

LeConte (1855, p. 430) holds that "Although very seldom seen except
when accidentally turned up by the spade or the plough, yet the Scaphiopus
is a very common animal in Georgia. Whenever the Southeast storms occur
at the Autumnal Equinox, the surface of the earth is covered with them, and
their dismal croaking adds to the horror of the howling winds and the deluges
of water which pour down from the heavens. They have also been found in
Connecticut and Massachusetts. When placed upon the ground they dig with
remarkable celerity, and soon entirely bury themselves in the earth."

Smith (1879, pp. 651, 652) observes: "For more than two years I have
been looking for the 'spade-foot' (Scaphiopus holbrookii) in and about New
Haven, confident that it occurred here and that careful search would reveal
it; but my efforts have been vain until very lately. Thursday, April 24th, I
saw some children gathered around an object on the pavement of Prospect
street, and I asked them what they had. They replied that they had dug up
a toad in the next yard. You can imagine my surprise and delight to behold
a real live 'spade-foot,' the first I had ever seen aUve. They wilhngly gave it
to me, and I carefully took it home with me and kept it alive in a large box
with plenty of earth and a tub of water.

"Tuesday morning (the 29th) I was shooting small birds near Fair Haven,
when I heard a most peculiar bellowing from a pond near by. I am more or
less famihar with all the ordinary sounds that come from a pond, and I jumped
at the conclusion that I heard the 'spade-foot.' On reaching the pond I saw
a sight I shall never forget ; the pond was rather small, and was swollen and
overflowing on account of the heavy rains of the two preceding days, and
swimming all over the surface, and at times uttering their peculiar bellow, were
forty or fifty of my long-sought friends. They would float or swim awkwardly

Plate XVIII

Spade-foot (Scaphiopus holbrookii)

1. Male and female, Chesser Id., Ga. August i8, 1922. Dorsal aspect, x 0.66.

2. Male and female (larger). Chesser Id., Ga. August 18, 1922. Ventral aspect.

X 0.66.

3. Adult, Chesser Id., Ga. August 18, 1922. Lateral aspect, x 0.8.

4. Male, Chesser Id., Ga. August 19, 1922. Lateral aspect, x 0.8.

5. Egg masses, Hilliard, Fla. August 17, 1922.

6. Eggs, stained, Chesser Id., Ga. Augu.st 18, 1922. x 1.45.

Scaphiopus holbrookii 79

along until they wished to favor me with a song, and then the accommodating
soloist would suddenly assume a perpendicular position as if a plummet had
been attached to his tail, his head alone swimming above the water, his white
throat dilated till it was three times the size of the head, his mouth closed
tight, he would sing his brief song and reassume his horizontal position. The
pond was quite deep in the middle, but I secured some specimens to prove my
statements on my return home.

"When I passed the pond again in the afternoon the same programme was
being carried out, but I could secure no more specimens. On my return home
I put my toad with the one I found Thursday, and in a few moments the male
(the last caught) had clasped the female very tightly and I was expecting to
raise some tadpoles, but they buried themselves in the earth the next day
without laying any eggs.

"In the afternoon of Tuesday a friend of mine, Mr. W. H. Fox, found the
Scaphiopus in a pond out on Prospect street, and secured quite a number of
specimens together with some spawn which he thinks belongs to this toad.

"The next day (Wednesday, April 30th) I visited my pond again with net
and pails, but the birds had flown without leaving a sign. Not a toad was to
be seen or heard, and no spawn but frog spawn could be found; but they may
have dropped it in the deeper water in the middle of the pond, out of my reach
and sight. Mr. Fox visited his pond also Wednesday, but could not find a
toad except the common one (Bufo americanus) .

"When I brought my first specimen home she buried herself in the earth,
but when I returned from Fair Haven she was swimming around in her tub of
water like the rest of them, and when I put the male in they stayed in the
water together. Wednesday morning when the toads in the two ponds had
disappeared, my pair had also buried themselves again in the earth in their
box. So I think I can judge of the movements of the free toads by watching the
movements of my captives."

Pike (1886, pp. 213-215) writes "Many years ago I had several in my garden
which became quite tame, and would allow me to take them in my hands.
They made circular holes in the ground about six inches deep, somewhat
turnip shaped. A few minutes sufficed for them to burrow out of sight. The
long feet, with the horny excrescence serving as an additional toe, and the
strong curved fingers enable the Spadefoot to make the excavation rapidly.
This is not by any means the completion of its home. The inside has to be
worked smoothly, and the earth prevented from falling in.

"This is done by the animal working its body with a circular motion, and
the operation would go on for an hour or more, and the liquid exuding from
its pores worked into the earth made it smooth, and formed a curious little
dwelling when completed. Round the top was a layer of viscious matter, and
woe betide any unwary insect that alighted on it. Closely concealed lay
Spadefoot, only the bright eyes visible, ever on the watch, and unerring in its
aim when any luckless fly intruded on the threshold. They appeared to be
greedy feeders, and I often amused myself by giving them insects, which they
seized with avidity as long as I supplied them.

8o Frogs of the Okefinokee Swamp

"This I find is the usual summer residence of the Spadefoot, and when
once domiciled, it rarely leaves home in the day-time. No two ever inhabit
the same hole, hence the name Hermit Spadefoot, or solitarius.

''When sharp summer rains fell they would quit their houses and seek
shelter under plants, but would not return to the holes they had left. As soon
as it was fine, about sunset or in the night, new houses were constructed with
great rapidity. In the fall, very heavy rains set in and one by one my pets
disappeared, generally in the night, and though I searched diligently for
them I could not find their hibernaculum, and presume they burrowed away
under the fence.

"Thirty-five years ago I exhibited one of these habitations, which was
made in a box sunk in my garden, with the animal in it, and read notes on it
which were published in the Proceedings of the Brooklyn Natural History
Society.

"The Scaphiopus changes much in color at various seasons. I have taken
this animal late in the fall when it might easily have passed by as only a dingy
young toad, but for the curious eyes which will always identify the Spade-
foot, no matter what its dress The irides are mottled gold and brown, and
are divided into four parts bj^ a notch at each quarter, giving a lozenge shape
to the large black pupil.

"Quite late in November, I dug one up from about a foot below the sur-
face of the earth, which was covered with decayed leaves. The head was
smooth dark brown, and the whole body a dingy dark olive, with faint lines
running from the eyes along the back, converging to a point at the rump.
Every part of the animal was tubercled, even to the eyelids, and the parotid
glands were greatly swollen. When first taken there was an orange tint over
the thighs and hands, but this soon faded in confinement.

"The little fellow took kindly to its imprisonment, grew very tame, and
looked quite comfortable in its large glass jar half filled with damp moss.
Sometimes when the moss seemed too wet I put in a lot quite dry. The cunn-
ing animal would look at it, toad-like, with its head on one side, and take in
the situation at once. It set to work and in a few minutes made a pretty little
arbor, quite thick behind but so thin in front that it could see through the
moss. It never appeared quite torpid, but only sound asleep at times, and
would wake up quickly if disturbed. A favorite position of my Spadefoot
was to crouch down flat with the hands turned in under the chin, the feet
turned up, and the long toes resting on the elbows.

"In April I took it out and found it as fat as when its winter rest began,
although it had not tasted any kind of food for over six months. * During this
long quiescence its coat had changed. The centre of the back was a bright
sandy color with a large dark irregular star edged with black. The whole
back and legs were heavily tubercled with a vivid red, chin white, abdomen
and inside of thighs a reddish purple."

* "In autumn the Scaphiopus feed voraciously and become very fat, and this seems to
keep them in good condition till spring again brings forth their insect food."

Scaphiopus holhrookii 8i

Hargitt (1888, pp. 535-537) writes: "A summer on Martha's Vineyard, and
the occasion of a sudden and tremendous rainfall, afforded the opportunity for
certain very interesting observations.

"My observations, as will be seen, add but little that is new; yet a record
of them may contribute somewhat to corroborate and extend that which does
exist.

"One afternoon, about August 10, 1887, while at work in the laboratory of
the Martha's Vineyard Summer Institute, in company with Dr. H. W. Conn,
Professor L. W. Chaney, and others, a very sudden and torrential rainstorm
occurred, lasting some two hours or more. During an interval of cessation
our attention was diverted by weird, plaintive sort of cries, which none at
first was able to explain. Darting out through the still-falling rain toward a
low sort of hollow, from which the cries seemed to come, it was found to have
been converted into quite a pond, though previously quite dry. In this, and
swimming about in a state of the greatest activity and excitement, were what
looked to be scores of toads. No difficulty was found in securing a few spec-
imens, which were at once identified as 'spadefoots.' Procuring a scoop-net
we took several dozens of them, leaving many more in the pond.

"This was about four o'clock in the afternoon, and they continued their
orgies till late at night. But I made careful examination the following morn-
ing, without finding a single specimen — not even a sign of one. In the water
I found plenty of the spawn attached to grass and floating in strings, loosely
attached to weeds — a fact which clearly indicated the purpose of their presence
and peculiar excitement.

"On account of the pressure of other studies, I was not able to watch the
development of the eggs. Indeed, I doubt whether they ever hatched, as the
pond was nearly dry before the close of the following day, and the soil, being
the loose sandy drift pecuhar to that locality, would not certainly retain
water for sufficeint time for the growth of the tadpole — if, indeed, for the
hatching of the eggs, though, as to this, last, I cannot say, as I left before it
could have occurred.

"This characteristic of the spawning habit is certainly peculair, and seems
somewhat difficult to explain. First, the lateness of the season is remarkable.
It is said that a related Eurpoean species breeds twice a year. Can it be
possible that such is the case with Scaphiopus? There are some facts which
seem to indicate that it might be, though it is hardly probable. Second, the
places of spawning is still more remarkable. From the observations of Dr.
Abbott and Colonel Pike, as well as my own, the choice seems to be for some
temporary sink-hole or surface-pond. If the conditions for development in
these places from speedy drainage, etc., were not so utterly precarious, it
might be thought a shrewd precaution for evading the natural enemies common
in the more permanent ponds and bogs. Altogether, the case seems to be quite
anomalous,

"But to refer again to the adults in the pond: There they were by scores.
Whence had they come, and in such numbers? In all probabilities, from the
ground of the bordering hillsides and environs. But not a single specimen was

82 Fi'ogs of the Okefinokee Swamp

seen out of the water, and that, too, notwithstanding we were at the pond
ahnost immediately following their first coming. If they had come from any
tolerable distance, it would seem that some late-comers would have been
detected. Again, their retreat must have been almost as sudden as their
appearance. I passed the pond about ten o'clock at night, and the air was
perfectly vocal with their never-to-be-forgotten notes. I went to the pond
early the following morning, but all was silent and deserted. Had they re-
turned to the ground? Such seems the most probable explanation. Yet so
carefully had they covered the retreat that not the slightest trace could be
found.

"Furthermore, their appearance itself seems to be capricious and phenom-
enal. I made inquiries of persons of observant habits as to any previous
occurrence in the vicinity, but was not able to find any account of them.

"It has been suggested that they are, doubtless, nocturnal in habit, and
that this explains, in a measure, their comparative rarity, even where known
to exist. I have no hesitancy in assenting to the nocturnal habit. It is quite
in keeping with the habit of many of the order; and the vertical pupil of the
eye points likewise to the same face. This, however, in itself, must be a com-
paratively small factor in the case, and, alone, would hardly insure it against
frequent detection any more than does a similar habit in many other nocturnal
animals. I had gone by this hollow repeatedly, night after night, both before
and after this appearance, and, though constantly on the alert to notice any-
thing of the sort, had no hint of their presence.

"Doubtless, the soHtary burrowing habit goes much further in explaining
its seclusion. But even this would be inadequate, unless the animal persis-
tently avoided all open and cultivated grounds. Such, only, would protect it
against frequent exposure by the spade or plow.

"Altogether, they are certainly the most peculiar and erratic of any of the
order; and, under the peculiar difficulties in the way of continuous study, it
will be long ere its life-history can be said to be thoroughly known. However,
the very difficulties add a charm to the investigation, which we may hope will
lead to success. To me, the brief research herein outlined has been full of the
liveliest interest, and, while but a mite toward the solution of the problem,
I shall hope that it may not be without some value when a final summary is
made."

Cope (1889, p. 301) maintains that, "This species, though so widely
distributed is seldom seen. After rains in spring and summer its cries may be
heard at night, proceeding always, so far as my experience goes, from tem-
porary pools. I have observed it twice in Pennsylvania, twice in New Jersey,
once in Massachusetts, on the main land opposite Martha's Vineyard. Speci-
mens from the latter locality which I kept in a vivarium buried themselves in
the earth by day, but issued at night-fall, and industriously explored their
surroundings. Their burrows were concealed by the loose earth which fell
into and filled them, but below this the bony top of the head could be always
found. Frequently one eye projected from the debris, presenting with its
brassy-colored iris a most singular appearance. On being irritated with a

Scaphiopus holbrookii 83

hard object they utter a clattering note entirely unlike that of the breeding
season."

In 1896 C. S. Brimley asserts, "I have occasionally dug them out of the
ground."

In 1899 G. A. Boulenger (pp. 790-793) in his paper describes the external
characters, measurements and skeleton in detail and remarks of its habits
thus: "The habits, so far as I have been able to observe them, are very similar
to those of Pelobates. They burrow in the soil in exactly the same manner and
come out only at night to feed. All my efforts to induce them to produce, when
irritated, the loud cries so striking in Pelohates have failed. On the contrary,
when teased, they assume a very humble appearance, bending down the head
at an angle to the vertebral column and shutting the eyes in a manner which is
well represented on the accompanying plate."

In 1904 C. C. Abbott (pp. 163, 164) holds the Spade-foots may be one
explanation of the reported showers of toads. He writes: "The frequent
references in newspapers to occurrences of 'showers of toads' have suggested
to the author that a condition in the life-history of the spade-foot toad, a
little-known and strictly nocturnal species, living in the ground, might explain
them more rationally than that the little batrachians are picked up by the
wind in one place and dropped in another, perhaps miles away, or that other
still more strange view quite common among the ignorant that toad-spawn
is sucked up by the sun and hatched in clouds, where the tadpoles remain
until they have advanced to the dignity of hoppers, when they fall to the
earth. Unlike the common toad and the frogs, the spade-foot toad (Scaph-
iopus solitarius) does not have a regular season for deposition of ova, but the
eggs may be laid at any time from April i to August 31. Furthermore, this
batrachian does not resort to permanent water-courses or ponds on such
errand, but takes advantage of temporary pools formed by showers of longer
duration than is usual. It is remarkable how admirably this strange irregular-
ity of an important event should be adapted to transitory conditions. Pools
of rainwater seldom remain long on the ground's surface. Soakage and evap-
oration soon obliterate them; but that this may not prove a fatal objection,
the eggs of the spade-foot toad hatch in about ninety-six hours, and in less
than two weeks, or fourteen days at most, the tadpole has become a terrestrial
animal or a 'hopper' and leaves its nursery. The development is even more
rapid occasionally, I am led to believe, being accelerated by excessive warmth
or retarded if the days are cool and cloudy.

"It will be readily seen that young spade-foot toads, congregated in or
immediately about a temporary pool, will not wander far from it when their
subterranean life begins, but will bury themselves in the comparatively moist
ground where they happen to be. Should, at this time of their limited wander-
ing, there occur one or more violent showers, the ground being wetted and
little pools formed, the young spade-foot toads would necessarily, we might
say, wander over a much wider extent of territory, and, escaping notice when
confined to one fast disappearing pool, would be observed when dotting the
ground over an extent perhaps of an acre or more. Seen thus, immediately

84 Frogs of the Okefinokee Swamp

after rain, and not previously noticed, the inference is not so strange that they
came to the earth with the rain, or that there had been a shower of toads
as well as of water."

In 1905 Fowler (1906, pp. 92, 93) writing of New Jersey forms gives, "the
following notes are on an example which I received from Mr. James A. G.
Rehn, during August of 1905. As it was brought alive from Palatka, Florida,
by Mr. Rehn I give it place only as a foot note. — This toad has a very peculair
habit of attempting to dig backwards by means of the black-edged spur on
the hind foot, which it will rub rather fast and with good force against the
palm if held in the hand. By a similar process it sinks into the ground or
among grass. The eyes can be depressed considerably and give the head quite
a different appearance if the animal is annoyed. It progressed usually by
short hops or leaps, though sometimes crawled or walked slowly a few steps.
In repose the usual position is to squat flat, and if then disturbed to inflate
the flanks greatly. When held in hand it uttered a note similar to the call of
Rana clamata, which may be said to resemble somewhat a smothered rattle
or trill of rather harsh and low tone. The throat is inflated, though not very
abnormally. This note seems to be due rather to discomfort than anything
else. In captivity I could not induce the specimen to eat."

W. T. Davis (1908, pp. 48, 49) gives the following observations on spade-
foots at Lakehurst, N. J., in 1906 and 1907. "During the heavy storms of
August I and 2, 1906, Mr. Louis H. Joutel observed many of these toads at
Lakehurst, N. J., in a depression temporarily filled with water. Their dis-
cordant notes could be heard a long distance. On July 20, 1907, I found at
Lakehurst two partly grown individuals that had fallen into a cemented
drain, up the steep sides of which they could not climb. On August 1 7 another
was found in the same drain, and still another at the bottom of a shallow well
that had become nearly dry. On October 12, 1907, with Professor Wm. M.
Wheeler and Mr. G. P. Engelhardt, we found a somewhat larger individual
in the same cement-lined drain. One of these toads was kept alive some time
and fed on flies, but died after being fed on green flies. Mr. Engelhardt also
had a frog die suddenly after it had devoured meat flies. The spade-foot toad
has also been found on Staten Island."

Dr. Frank Overton (1914, pp. 29, 30) who unselfishly introduced the frogs
and toads of Long Island to the naturalists of Greater New York, just as
earlier W. T. Davis did the same for some of the rarer species of New Jersey,
has in recent years given more attention to spadefoots than any naturaUst in
the North. From him we cull excerpts under several topics. He writes thus:
"Spadefoot toads are considered to be rare, and few detailed observations of
them have been made. Dozens of them suddenly appeared in a temporary
pool near the Bay Avenue school building in Patchogue during the first week
of April, 191 2, after a series of hard rains. April seventh was warm and pleas-
ant, and the toads were noisy all day. The temperature fell below freezing in
the evening, and all the toads disappeared until the sixteenth, when the air
again became warm, and a number reappeared from that day only. The tem-
perature again fell in the evening, and no more spadefoots were heard until

Scaphiopus holbrookii 85

the twenty-seventh, when a few were heard after a hard rain. None were
heard or seen after that date. A great number of eggs were laid, but no tad-
poles survived the frosts.

"The spadefoot toads suddenly reappeared on the evening of April 12,

19 13, in several pools near Bay Avenue after a warm, hard rain. They were
noisy on the thirteenth, but after that date the weather was cold and only a
few were seen until the twenty-third. Then about fifty appeared in the pool
which had just refilled on the site of the pool in which they were seen in 191 2.
After two or three days all the toads had disappeared. Hosts of tadpoles were
hatched but most of them were destroyed on account of their pools being
filled in with soil. Those that survived completed their transformation into
fully formed toads by the middle of June.

"A few spadefoot toads appeared in a small permanent pool near the Bay
Avenue school on April 27 and 28, 1914, and laid considerable amount of
spawn. There was a prolonged rain on May fourth and fifth and on the
evening of the fifth great numbers of the toads appeared in the pool. About
one hundred pairs were mated in the pool all through the following day, and
their method of laying eggs was readily observed. The chorus of unmated
males was loud on the evening of the sixth, but not one of the toads could be
found on the seventh, although the temperature continued warm. On July 7,

1 9 14, numbers of spadefoot toads appeared on the salt marshes adjoining the
shore at Bay avenue, Patchogue, after a week of rainy days. They sang in a
loud chorus and many were mated. None could be found on the next day."
"The tadpoles of spadefoot toads have been found in Oakdale and Speonk.
The toads are widely distributed, and their times of appearance are as regular
as those of other toads. They escape observation on account of their burrow-
ing habits, and their short stay in the breeding pools. I made sure of finding
them by hiring a small boy to visit the locality every evening." In many
respects the author agrees with the last paragraph from observations made
on Scaphiopus couchi and Scaphiopus hammondii. It is an interesting fact
that Prof. R. J. Gilmore of Colorado College, Colorado Springs, Colo.,
regularly and yearly uses Scaphiopus for his class use in biological courses
instead of Bufo or Rana species.

After the pubhcation of his Long Island Frogs and Toads, Overton, 191 5,
pp. 17, 52, 53, followed the yearly appearance of spade-foots with two notes in
Copeia. They are: "vSpadefoot toads {Scaphiopus holbrookii) appeared in
Patchogue on the evening of May 22, 1915, for at least the fourth successive
year. The first hard rain of spring occurred on the night of May 21, and on
the evening of May 22, great numbers of toads were present in the same pool
in which they had been seen in previous years. On the morning of the 23rd
nearly every blade of grass in the pool was covered with their eggs, and by
night not a toad remained in sight. The sudden emergence of the toads from
their underground retreats and their extremely short stay in breeding pools,
probably accounts for the former belief that years usually elapse between
their appearances."

86 Frogs of the Okefinokee Swamp

Later that year he gave another note (19 15, pp. 52, 53) : "On the evening
of Aug. 4, 191 5, enormous numbers of Spadefoot Toads {Scaphioyus hol-
brookii) appeared in Patchogue, Yaphank, Middle Island and Coram. I made
a twenty-five mile circuit through the villages named, and found the toads in
practically every pool. There had been an extremely hard rain during the two
previous days."

The next year on Long Island J. T. Nichols (1916, pp. 59, 60) makes the
following observations: "The morning of June 17, 19 16, after heavy warm rain
the preceding night there were singing Spadefoot Toads in a woodland pool
beside a road. At mid-day one was seen to cross the road and hop away from
the pool into the woods. Its color matched the leaf-carpet wonderfully.
Others remained all day.

"One was captured in the pool and liberated the following day. It is
remarkable how completely it was able to hide in a closely cut green lawn in
bright sunlight by crouching at the bases of the grass. When liberated in the
woods it disappeared backwards under the leaves, and remained with just
the nose showing at the bottom of its entrance. On June 25, it was found
again in this same spot under the fallen leaves in a shallow burrow in the
ground, its nose showing. When disturbed it turned sideways, thus with-
drawing completely and filling the mouth of the depression with sandy soil.
July 3, on scraping away the dead leaves, there was no sign of the toad, but a
spot of loose soil detected was investigated disclosing it at a depth of about
I 1/2 inches. This was the last seen of that particular individual, as on July
9, there remained only a neat steeply-slanting burrow, about 3 inches deep,
empty.

"A steady rain commencing the night before, continued through July 23,
on which afternoon Spad'efoot Toads were singing in a pool in pasture land
near stands of trees. During a temporary silence cattle came close to the pool,
only to gallop away in alarm when the noise recommenced. Investigation
disclosed singing Spadefoot s also in the woodland pool occupied several weeks
earlier, and a greater number in woods now flooded just across the road.

"Points of interest in these data are coloration in the woods, skill at hiding,
recurrence in the same pool with favorable conditions after 36 days (See
Overton, Copeia, Nos. 20 and 24), and an individual's remaining 15 days in
one spot just under the fallen woodland leaves."

VOICE

Andrew Nichols (1852, pp. 113-115) gives considerable on the voice and
habits of this species.

"In a shallow basin surrounded by ledges of green stone rock, which retains
water during the winter and spring, and is occasionally filled in summer by
great rains to the depth of one to four feet, on the brow of a hill in Danvers, over
which the old Essex Turnpike crosses, and near the intersection of this road
by the Newburyport Turnpike, an interesting colony of this rare reptile,
hitherto unobserved north of South Carolina, has been lately discovered.
Somewhere about the years 1810, 1811 or 1812, subsequent to a great rain in

Scaphiopus holhrookii 87

summer several frogs of this curious species were noticed by John Swinnerton
Esq., now deceased, who resided very near this spot for sixty successive years,
and had ample opportunity to observe them. Their numbers in this, their
first appearance, were as great as any time since, determined by the testimony
of several witnesses. At this time, during one or two days and nights, they
were very noisy and actively engaged in fulfilling the great fiat of creation;
and soon afterward they disappeared. Nothing more was seen of them for
several years. Their voices or note so nearly resembled that of young crows
that it attracted the attention of Mr. Elijah Pope, a farmer, who lived half a
mile distant; and, accompanied by his son, sallied forth, gun in hand, to kill
what he so naturally imagined was mischievous birds in his corn-field. From
this Httle anecdote I am enabled to determine their first observed appearance
with some degree of certainty; thus defining the time as about the first of
September, when corn is in a state to be fed upon by a crow: while, again,
the year mentioned above, agree with a statement of the son, Mr. N. Pope,
who from his own present age, concludes that 1812 or a year or two earlier,
was the time of their first visit.

"So far as recollection serves, nothing more was seen of them until July
1825, on a day memorable for the passage of La Fayette over the Newbury-
port Turnpike close by, on his return to Boston from his eastern tour; at
which time their voices added to the welcome greetings of a nation's guest!
Since this last date, thus rendered conspicuous as well as certain, whenever
the basin has been filled in warm weather, these reptiles invariably make
their appearance. This has occurred, however, only three times, viz., August
12th, 1834; again in the summer of a year whose date is forgotten; and on
June 1 6th, 1842. The forenoon of this day, last mentioned, was dark and
rainy, as the day and night previous had been. Their croaking attracted the
attention of an acquaintance of mine, and information of their appearance,
with a pair of the frogs, was forwarded to me by Mr. Amos Swinnerton."

In 1884 both Dr. Charles C. Abbott in New Jersey and Col. Nicholas Pike
observed this species in congress. The former reports on voice as follows:

'Tn a sink-hole in a dry upland field near Trenton, New Jersey, on April
10, 1884, there suddenly appeared a large colony of hermit spade-foot toads
(Scaphiopus holhrookii), which, by their remarkable cries, attracted the atten-
tion of every one passing by. So unlike the cries of any other of our batrach-
ians were their utterances, that all who heard them were attracted to the spot,
and wondered, when they saw the animals, that so great a volume of sound
could issue from so small an animal. One need not wonder, however, on this
point if he will but examine the development of the animal's vocal cords.The
machinery for producing sounds equal to an ordinary steam whistle are
apparently contained in the throat of this rare and curious batrachian.
Holbrook, in his diagnosis of the genus Scaphiopus, refers to the 'sub-gular
vocal sac' of the males; but it must be inferred that the famales are voiceless.
That they are not so noisy is probable, but occasion requiring, they can readily
make themselves heard.

88 Frogs of the Okefinokee Swamp

"It was not long, however, before the spade-foots again became the
prominent figure of the fauna of the neighborhood. During the night of June
25-26, a violent north-east storm arose, and rain fell in torrents. The sink-
hole, which for weeks had been nearly dry, was again flooded, and on the
afternoon of the 26th was literally alive with these rare toads. Sitting upon
every projecting stick or tuft of grass, or swimming with their heads above
the surface of the water, were spade-foots by the hundred, and every one ap-
parently uttering those shrill, ear-piercing groans that only these batrachians
can utter. Not only during the day but all night their cries were kept up.
The following day there was no abatement, but during the night the sound
decreased. On the morning of the 28th not an individual was to be seen or
heard.

"I have already referred to the wonderful noises made by these animals
when they congregate in pools for the purpose of spawning. At no other time
do they appear to be vocal, and the question naturally arises, why, when the
animal leads a life that requires no such power except for two or three days in
a year, should its utterances be far louder than any or all the frogs and toads
of the same locality combined? Although the animal is strictly crepuscular
and not diurnal, it could readily find a mate guided by sight, and the purpose
of the deafening epithalamium is somewhat hard to determine. If it could be
shown that they call to each other from far distant points, the difficulty would
disappear, but this they are not known to do. Apparently it is not until they
are congregated in some available pool that they sing, if singing it can be
called. No words yet in use in our language can fairly describe their utter-
ances, which, it may be presumed, are expressions of dehght at meeting."

To Brimley (1896, p. 501) "The cry was not much louder than that of the
common toad." To Ditmars (1905, p. 191) "the voice of the male resembles
the tremulous call of the common toad, but is slightly louder."

Fowler (1907, pp. 96, 97) writes: "They are noted for their irregular and
erratic appearance during warm weather, their burrowing in the ground
tending to their escaping observation. About Trenton, according to Dr.
Abbott, it may appear at any time between May and September, and erratical-
ly in abundance. They make a great noise, and as the egg-laying varies in
time, their notes may also be expected to be heard at different times. During
copulation, both animals roar so that they may be heard at the distance of
half a mile. Their roar is not like that of the common toad, Bufo, and their
vocal apparatus is not especially different during the operation. They will
also roar at other times, as when disturbed or during heavy rains. Mr. C. C.
Abbott informs me that he heard a few during the spring of 1904 and 1906.
Dr. Dahlgren tells me that he secured examples near Trenton from the same
locality where Dr. Abbott made his observations."

Dr. Overton (1914, pp. 28, 29) writes of its voice thus: "A mud puddle, or
a temporary pool formed by a prolonged rain in early April sometimes swarms
with toads that groan and squawk in a most unpleasant manner. Each squawk
is hke the groan made by a deep-voiced man having a tooth pulled. It may
also be compared to the squawk made by a big rooster caught in the night.

Scaphiopus holhrookii 89

Such a sound coming from a temporary pool in April is almost surely the voice
of a spadefoot toad.

"A spadefoot toad makes its noise while it lies sprawled on the surface of
the water. When it begins to sing, it suddenly distends its throat into a white
pouch that is about double the diameter of its head. The result is that the
pouch, floating on the water, Hfts the toad's head up suddenly while the hinder
parts of the body sink beneath the surface. At the expiration of the sound,
the toad bobs back to its sprawling position. Each vocal effort lasts about a
second. An active toad will repeat the sound about every ten seconds. A
spadefoot singing in a pool on a dark night may be recognized by its vocal
sac which appears like a big white bubble in the light of the lantern."

Shortly after Dr. Dunn's hst of Reptiles and Amphibia of Virginia ap-
peared H. W. Fowler adds Scaphiopus holbrookii to the list with these notes:
"On July 15, 1906, Dr. Henry Fox obtained a fine example of Scaphiopus
holbrookii which he forwarded soon after. He wrote: "The specimen as well
as several other examples were exposed in plowing a stubble-field and attracted
attention by the extraordinary loud and shrill cry they made when crushed by
the plow. Mr. E. R. Dunn informs me this amphibian has not been definitely
recorded from Virginia and the above examples all observed at Tappahannuk,
are, therefore, of interest. In this connection I may also mention that under
date of June 7, 19 16, Mr. H. Walker Hand writes from Cape May, N. J. 'On
May 28, Dr. J. S. Eldridge and myself were walking along a wood-road and
heard a peculiar cry coming from the ground. I had heard the same before,
but could not find the source. This time I dug down and the sound continued
even as I was digging. At a depth of about three inches a spadefoot was
found snugly fixed. He never stopped calling until I lifted him out.' "

In 1920 from Dade County, Florida, Deckert reports that "During a
prolonged thunder-storm many of the Spadefoot Toads were encountered by
the writer in the streets south of the Miami River on the afternoon of May 16,
and during the night were found breeding at 19th Street and Avenue H., also
at 22nd Street and Miami Avenue, and great numbers were reported from
the low grounds near the 'Alligator farm/ Miami. Their cries sound hke 'Ow,
Ow,' and 'Miow,' but the latter much deeper in tone than the well-known cat
cry. The noise made by a dozen males is deafening when one is near, though
the call lacks carrying power."

On August 16, 1928, about one mile southeast of Hilliard, Florida, we were
slowly travelling along the Dixie Highway detour. W^e were helping tourists
over an overflowed area when Mr. Francis Harper espied a curious hawk and
started after it. The search led him across the Dixie Highway and he heard
what he at first mistook for young crows. (In this connection it is interesting
to recall Mr. Andrew Nichols' account wherein spade-foot calls were Hkened to
the call of young crows) they proved to be spade-foots. We, therefore, left
the detour and went through the woods to camp beside the five or six ponds
where the congress was. We could hear them for about a half mile's distance.
Nearby it sounds somewhat like where where, where, where and so on. Once one
of our party said it sounded at a distance like the calling or snarling of a cross

Qo Frogs of the Okefinokee Swamp

baby. To the writer at first they sounded Hke some Rana in distress, much
as they call when a snake or turtle have them. At a distance at other times
they sounded like young herons in a heronry. They may make as much noise
as a steam calliope but it hardly sounds hke it. Mr. F. Harper characterizes
the call as "naarh, naarh — complaining, nasal, not shrill or high pitched."

When we first approached the males called on all sides in full daylight,
almost at our very feet. The throat by day is a beautiful glistening white
when the male is croaking. The throat when inflated may be a ball from i-
I 1/2 inches in diameter. The male before he croaks lays on the water's
surface with hind legs partially submerged. When he croaks he tips the hind
end of the body and hind legs beneath the water and the head is reared to a
45°-75° angle with the water's surface. When at the height of the performance
or slightly before he closes his eyes. Then the throat deflates and body
inflates. It croaks about once in every two seconds.

Some occasionally question whether female frogs croak or assert they do
not. Often a female when picked up will give a croak or squawk not like the
breeding call of the male. Sometimes when two or more males are struggling
for a female she will squawk, possibly because it may be almost too much for
her to bear. Normally the male and female of a mated pair seldom croak.
It is usually the males which are searching a mate or annoying the already
mated pairs.

MATING

Male {From life, Aug. 18, 1922). Stripe from eye back to vent lemon
yellow or greenish yellow. Snout mummy brown or prout's brown. Other
light colors of the back are same as stripe from eye to vent, i. e., lemon yellow
or greenish yellow, so also on the sides. Snout sometimes blackish brown (i)
or bone brown. This is also the remaining color of the back. Sometimes the
back color is virtually black. Limbs on dorsum much the same color or
slightly washed out more greenish yellow than back light colors. Under parts
of hind limbs, lower belly light grayish vinaceous. Rest of under parts white
especially the white glistening throat when croaking is done. First three
fingers with black excrescences. Space of hind foot dark-edged and tip of ist
toe. Web much darker than grayish vinaceous web of female. Male have
wider and larger hind feet. Iris light greenish yellow with black on outer
rim. Pupil vertical.

Female {Frofu life, Aug. 18, 1922). Sometimes uniform warm sepia or bone
brown above. Throat and breast white. Underside of forelimbs and hind
hmbs and lower belly light grayish vinaceous. Sometimes the stripe on fe-
males may be sulphur yellow instead of the intense greenish yellow of male.

Structural differences. The first character recorded is "Males with a sub-
gular vocal sac (Holbrook 1842, p. 109). Dumeril and Bibron (1841, Vol.
VIII, p. 472) write that "The males under the throat have an internal vocal
vesicle which communicates with the mouth by two large longitudinal slits,
placed one to the right and one to the left of the tongue." Or "Un sac vocal
sous-gulaire, chesles males." (Same p. 471).

Scaphiopus holhrookii 91

In the field we made the following notes:

(a) Strangely enough the croaking males were not so hard to approach
in the daytime as at night when we tried to photograph them. At night we
found them more wild and shy, restless. They proved harder to see and hold
by flashlight than many species of toads or frogs. Often when croaking they
were just out of the range of the flashhght.

(b) Almost always each male of a pair had a predominance of yellowish
hues and the female inclines to the brownish hues.

(c) The throat when inflated glistened by day or under the flashlight in
the night.

(d) Apparently the females seem to have hind feet very perceptibly
smaller and lighter than in the male.

(e) "The male seemed to have the rear end of the body more of an
extended ridge than in axillary, pectoral embracing forms."

(f) Often a male if placed on his back would draw up his hind legs and
fore legs and rest on his back as a gopher frog will on his haunches if leaned
up against some object.

(g) Males have the inner edges of first two fingers and sometimes the
third finger with excrescences; and the first two fingers and the fingers in
general are broader, less slender than in the females.

(h) The males and females are not so dissimilar in size as in some species.
The males externally are visible as such at about 53 or 54 mm., the females at
50 or 51 mm. Presumably, according to our presumptive evidence of growth,
these are 4-year-olds.

Duration, day or night. Pike (1886, p. 218) says "the Spadefoot has a
great dislike to water, and when forced to it for breeding purposes does not
remain in it long, from three to five days at most. The embrace often takes
place on land, as it does occasionally in toads, so that they only enter the
water for the purpose of spawning. Their wonderful screaming chorus is
kept up the whole time the animals are in coitu, and is the love song of the
males, the females having only a low guttural grunt. As soon as possible the
sexes separate and seek their summer homes, where they lead solitary lives
till they have to seek their winter's retreat." In 1920 we wrote that "The
hermit spadefoot toad appears suddenly after prolonged rains in April and
May or sometimes June or July. At the breeding season it is fond of sprawl-
ing out on the surface of the water as a wood frog does, and it is from this
position that it croaks. This species gathers in large breeding assemblies like
toads, and the matings are as spirited."

On August 16, 1928, beyond noon we found these spade-foots in congress.
They continued all that afternoon and through the night, but early the next
morning, August 17, not a one was to be found in pond nor nearby. All had
left the pond or were in hiding in the pond or most presumably had burrow-
ed in the nearby soil. If heavy rainy weather is on they like other species of
spadefoots will breed by day, even start first in the daytime but darkness
is the preferred period for spade-foots in general (.S. holhrookii, S. couchi
and >S. hammondii) .

92 Frogs of the Okefinokee Swamp

The males in ardor are the equal of any toad. The tumbles and turmoils
are just as numerous as in a toad congress.

Amplexation (Normal, abnormal, cross). Brimley (1896, p. 501) in ob-
serving fifty breeding frogs writes that "In every case the grasp of the male
was inguinal." Richard Deckert's Fig. i in Overton's Long Island Frogs and
Toads portrays an inguinal embrace, in all three pairs pictured. In 1920
(p. 22) we say that "The male seizes the female just ahead of the hind legs
(inguinal fashion), a form of embrace not known in any other American form
except in the narrow-mouthed toads . . . ." How the last slip on narrow-
mouthed toads happened is beyond us now. Gastrophryne texense and Gas-
trophryne carolinensis embrace axillary fashion.

Normally a male embraces in inguinal fashion. In the five or six ponds
we saw no other fashion for ovulating pairs. Even when fertilizing the eggs
the male keeps the inguinal fashion but he draws up his vent near that of the
female. A male of a mated pair can be taken away and another substituted.
Almost invariably the second instantly remains on the female. Often one
will see a male rush for another male or for a mated pair. Occasionally one
sees a female with a mated male and another male atop the mating one.
When spade-foots are actually ovulating the male usually closes his eyes.

Several abnormal embraces were observed. These are usually preliminary
and not of ovulation period. One female was held by a male which had its
left arm above and behind the foreleg of the female and male's right arm was
just ahead of the right hind leg. Another male held her from below, his left
arm behind her right arm and his right arm around her left side of waist. At
times both of his arms were just in front of the hind legs.

In the ponds were narrow-mouthed toads {Gastrophryne) pine woods tree
frogs ( Hyla femoralis) cricket-frogs (Acris gryllus) and Florida tree frogs ( Hyla
gratiosa) breeding in great profusion. None were large enough for cross
embraces except with the last. In one pond we found a male spadefoot 62
mm. long embracing a ripe female Hyla gratiosa 58 mm. long. In the ponds
were several normal Hyla gratiosa pairs.

OVULATION

Habitat. The congress of ovulation we found Aug. 16 and 17, 1928 in
cut over pine barrens in shallow depressions the water in one pond being i-
I 1/2 inches deep, in several 4-6 inches, in another 6-8 inches, and in a last
pond 12-18 inches deep. The pools were in what might be called moist pine
barrens or it might be an old field, a cypress pond not being far away. One
pond was a depression in a wood road. Considerable of Rhexia, Hypericum,
sedges and grasses were in the five or six ponds. In the ponds were numerous
Acris gryllus, Gastrophryne carolinensis, Hyla femoralis and Hyla gratiosa
breeding. All in all the pools were indeed very temporary.

Period. In South Carolina Holbrook (1842, p. 1 1 1) found that 'Tt appears
early in March, after the first heavy rains of spring, and at once seeks its
mate."

Scaphiopus holhrookii 93

Pike (1886, p. 220) writes: "In the pond where I found them in 1884 there
was abundant water in 1885, fed by springs, and to my eye the same condi-
tions obtained this year. It is evident Spadefoot thought differently, and
resorted to a pool a quarter of a mile away, for some reasons unknown to me.
I believe they breed ewery year, but change their locahty, as the winters,
though often severe on Long Island, certainly do not diminish the numbers
of the Scaphiopus.'^

In Raleigh, North Carolina, Brimley (1896, p. 59) recorded "Last May I
collected fifty breeding in a pool only a few yards from my house." In Florida,
T. VanHyning (1923, p. 68) writes: "On the night of March 18, while some
of the boys of the biological class of the University of Florida, Gainesville,
Florida, were collecting frogs, among other species taken were thirty specimens
of the Florida Tree Frog, Hyla gratiosa LeConte ; eight specimens of the Goph-
er-Frog, Rana aesopus (Cope), and two of the Spadefoot Toad Scaphiopus
holhrookii (Harlan).

"These and other species were all in a pond breeding near the University.
One of the party collected eggs, and brought living specimens which have
spawned since . . . ."

The summation of 23 congresses from 1910 to 1922 give two for March,
four for April, five for May, three for June, three for July, three for August
and two for September, or eleven for the spring months, nine for the summer
months, and three for September, the modal month being May. The two
March records are: the first for South Carolina and Georgia; and the second
for Florida.

Our only record of ovulation observed is for August 16, 17, 1928.

Temperature and humidity. On August 16 and 17, 1928, when we found
the spadefoot congress we had no thermometers with us. We record that the
water must have been 75° to 80° when the rain was on and much higher when
the sun came out. The rain was so heavy that many cars were stuck on the
detour ahead of us and streams were flowing over the wooded bridges.

For five stations around the swamp for the two days, August 16 and 17,
the air minima ranged from 69°-74°, with an average of 71° and the maxima
ranged from 82°-9i°, with average of 87°. These stations average .70 inches
of rain, e. g.. Glen St. Mary, Florida, had 1.80 inches August 16 and a trace
the next day; Waycross had .69 inches August 16 and .55 inches August 17.
But Hilliard where we were had .56 inches August 16 and 3.47 inches August
17 or 4.03 inches when the congress was in progress.

Egg-laying process. Overton (19 14, p. 29) describes it thus: "The only
time when spadefoot toads are not shy is while they are actually laying thier
eggs. A pair of toads about to deposit a mass of eggs will cling to a stiff spear
of grass about a foot beneath the surface of the water and will slowly crawl up
the stem, depositing a string of about two hundred eggs enclosed in a gelatin-
ous envelope about as large around as the toad's leg. Each batch will be laid
within five minutes, and during that time the pair may be approached readily.
A toad will lay its eggs at intervals within a very few hours, and will then
disappear from the pool." Deckert's drawing (Fig. i, p. 39), May 12, 1914,

94 Frogs of the Okefinokee Swamp

in this work is a very good portrayal of the process and in accord with our
experiences in northern Florida.

One of our short field notes is as follows: "Spade-foots lay eggs on twigs
(upright). Water 6 inches or more deep in first pond. Pair lays a band on a
twig in about a minute or less. Female seizes the twig with forelegs and
climbs branches with forelegs and hindlegs. Male embraces inguinal fashion.

"Must be 50 pairs or more in one pond which is 4 or 5 inches deep. The
pair leave the surface and start for the bottom and the female moves up the
stem toward the surface to the end of the twig. Sometimes horizontal posi-
tion in laying, usually vertical. Intervals of laying. One pair has 15 seconds
laying. Rested one minute. Laid. Rested 3 minutes. Ten seconds for
laying. . . .

Often a female will hold her head downward at first. From time to time
the female will cross fore feet and hind feet as she climbs a stem."

To make sure our lone opportunity for observation on the breeding of the
Spade-foots in 192 1 and 1922 was used to its utmost we asked Mrs. Wright
to record independently her own findings. Under egg laying we presented
her notes : "We started from Callahan in a hard rain, a Uttle before noon. On
a detour two miles south of Hilliard, we stopped for cars going across a swollen
creek. Francis went to look for birds and heard Spade-foots calling. We drove
through the woods and back on the Dixie Highway and pitched camp, one
mile south of Hilliard on an oak ridge. An old road filled with water made a
shallow pond, and here we saw the males croaking, their white throats looking
like shiny white golf balls. Just beyond was a shallow surface pool, caused
by the heavy rain. The ground was covered with herbs; a little Xyris, a few
sedges, Rhexias, small umbelhferous plant with violet shaped leaf, wire grass
and a few Hypericums. The Spade-foots were calling here, and in another
similar pool and in a third deeper pool as well. At a distance the chorus
sounded like young crows trying to call. The pond was filled with mated
pairs.

"A few eggs had been laid. The eggs were laid in more or less irregular
band form along the grass blades or plant stems. In the third pond where
the water was deep, the bands were long. The pair might be floating on the
surface. When ready to lay, they went to the bottom of the pond, often the
male with his eyes closed, and the female with her's partly closed. They
moved around slowly on the bottom, or rested a minute. When she found a
stem to suit her, she seized it with her front feet and pushed with her hind
feet. The male clung close to her back, his chin tight against her back. (One
we photographed had an abrazed chin as if from pressure). He held his knees
against her knees, or sometimes his feet which are conspicuously broad, were
pressed against her feet. She walked or climbed up the grass blade or along
it, if it fell to horizontal position, pressed vent against the blade as she laid
the eggs. He humped his back to press his vent close to her's while she was
laying. As they reached the top of the blade, they sometimes moved imme-
diately to a nearby one, or rested a short period. When first laid the eggs had
a irregular band appearance as they were strung along the blade, sometimes

Plate XIX

Spade-foot (Scaphiopus holbrookii)

1 . Rain filled depression in the pine barrens, Hilliard, Fla. August 17, 1922.

2. 3. 4. Males croaking, Hilliard, Fla. August 16, 1922. Flashlights.

5. Eggs, HiUiard, Fla. August 16, 1922.

6. 7. Tadpoles, Camp Pinckney, Ga. June 22, 1922.

8. 9. ID. II . Tadpole and transforming frogs, Chesser Id., Ga. June, 28 1922.

to

Scaphiopus holbrookii 95

being much thicker as if more eggs had been emitted at such periods. When
first laid, they had a brownish appearance with conspicuous creamy-white
vegetative pole. As the jelly swelled and the eggs all turned right side up,
they looked very black.

"By the next morning, some eggs were almost ready to hatch (these must
have been the ones we found when we first found the pond). The other clusters
were swollen into loose, irregular, elongate bunches attached to the stems
which were tipped so that many times the bunches lay lengthwise on the water.
There seemed to be a tendency for the bunches of eggs to be more or less
clustered in areas. We noticed many pairs close together that first afternoon,
and unattached males trying to get at a female making tangled masses of
toads. There was a strong chorus that night and by the next morning the
pond was all churned up and muddy. Many, many eggs were there, but no
toads. The story was told for the season."

EGGS

Attachment, egg mass, egg description. On May 4, 1863 Mr. F. W. Putnam
(1863, p. 229) made some statements concerning the frogs and toads found
about Cambridge, Mass.

"The Scaphiopus solitarius Holbrook appeared in their old place, near the
Botanical Gardens, in large numbers on the 19th of April, and commenced
the same day to lay their eggs in bunches of about one, two and three inches
in diameter; these bunches were floating on the water and were not attached
to the grass, as was the case when seen in the previous years. On the 29th
of April, another set of Scaphiopus visited the place, and laid eggs, which
were attached to the grass as formerly. The tadpoles of this species are
hatched in about six days."

Pike (1886, pp. 218, 219) compares these eggs and toads eggs. "On the
ridge extending from East New York to Jamaica, one of the most elevated
parts of Long Island, there are ten or twelve ponds, some fed by springs and
constant, others often filled by winter's snows and rains. This spring I
worked them all over with my net, and though I heard no screeching, yet as
the Scaphiopus is far from rare on the hills near by, I felt sure they must
breed in some of these ponds. Toads, I know, also swarm in the vicinity, and
on the 17th of July I fished up what I took to be toad spawn, although not in
chaplets, and only slightly attached to some weeds floating about in the
water. The eggs were evidently laid only a few days before."

In 1897 Wm. L. Sherwood (1898, p. 18) holds that "The eggs are laid any
time from April to June in bunches from one to three inches in diameter, and
are placed around a spike of grass."

In 1899 G. A. Boulenger speaks of their eggs and breeding thus: "I had
appHed last summer to Messrs. Brimley, in North Carolina, where the Spade-
foot is abundant, who kindly informed me that the eggs are laid early in
spring, in strings resembling those of toads, but thicker and with the vitelline
spheres more irregularly disposed — in fact, as I infer, not unlike those of
Pelobates. They added that the season was then too far advanced for tadpoles

g6 Frogs of the Okefinokee Swamp

to be procured, as their development is comparatively rapid, and the pools in
which they are reared dry up by the end of spring. I have therefore to post-
pone a description of the tadpole, which I hope, however, to supply ere long."

Ditmars (1905, p. 191) holds "The eggs are similar to those of the toad,
and are laid in strings." By no stretch of description could one hold them the
same as those of the common toad.

(See Overton's notes under egg laying process and Deckert's Fig. i for the
appearance of eggs at or immediately after ovulation).

In 1920 (p. 27) before our own first-hand experience with spade-foot eggs
we repeated that "the hermit spadefoot lays eggs in bands like the European
forms of this family, . . . ." But it should be said that soon these bands
appear as cylinders. In 1923 (a p. 406) we speak of the "eggs in bands, later
cylinders." Or 1923 (b p. 34) we hold that "from April isth or earlier to

Sept. I St., six forms breed Scaphiopus holbrookii lays bands of eggs

which soon become cylinders." In 1924 (pp. 377, 379) we record the "egg
mass (as) submerged. Eggs laid in bands which soon become loose cylinders
extending along plant stems or grass blades; vitellus 1.4 to 2.0 mm.; envelope
3.8-5.6 mm. Egg complement 2,332. Season Apr. 15 or earlier to August
17." "These (early species) are usually with firm jelly envelopes except for
Pseudacris triseriata and Scaphiopus holbrookii, which have the consistency
intermediate between the firm jellies of early breeders and the loose surface
films of later breeders."

In one pond in an area 4 feet square were 1 2 pairs. Before a congress is
over the stems and vegetation at times gets tied up in a tangle with the inter-
locking and heavy egg masses. Some stems are with egg masses bent over
and such horizontal stems are often connected to a vertical one by other jelly
masses. A few fresh laid bands were as follows: i inch long, 2 inches, 4
inches. When the bands expand the band may be i 1/3-1 3/4 inches in
diameter. Usually the bands were about as long as the pond was deep. Most
were, therefore, 1-6 inches long. But in the one deep pond were bands very
much longer, 8-12 inches. In other words the female goes to the bottom of
pond and crawls up to or almost to the surface before she ceases ovulation.

The next morning at 10:00-10:30 a.m. the spade-foots having left before
daybreak, in an area 6-8 feet in diameter every grass blade was covered and
bent over from the weight of one or more masses. The masses were swollen
(not now band-Hke) and some almost hatched. Masses around the stems
were i inch wide or sometimes i 1/2-2 inches in width. If the original band
be short, the final swollen mass may at times look squarish at the surface or
be cuboid in reality. Most of the masses looked like cylinders of jelly or
round masses. The pond was just packed with these masses and the water
had a milky appearance.

We made several field observations and measurements on the individual
eggs. Mrs. Wright made these three determinations: In one lot, the vitelli
were 1.6 mm., 1.6, 1.8 mm., and envelopes 4.8 mm., 4.8mm., 5-2-5-0 mm., and
in mass the jelly more or less merged. Another set had measurements: 1.4

Scaphiopus holhrookii 97

mm., 1.6, 1.6, 1.6, 1.8 mm. for vitelli and 3.8,4.0,4.0,4.0, 4.5 for envelopes.
A third group had 1.4, 1.6, 1.6 mm. for viteHi and 4.0, 4.0, 4.2 mm. for
envelopes.

On three other lots the writer secured the following: One set give 1.4, 1.6,
1.6, 1.7, 1.8 mm. for vitelH and 4.2, 4.6, 5.4, 5.5, 5.6 mm. for envelopes.
Another group where egg envelope a day old or not fresh, tended to be ellip-
tical, we had vitelli 1.5, 1.7, 1.7, 1.7, 1.7, 2.0 mm. and envelopes 4.0, 4.1, 4.4,
4.5, 4.5 mm. The third group were one day or a little less old when pre-
served and jelly was more or less merged or there was no regularity of envelope
outline at all. At times an egg would hang at the end of a jelly cylinder as if
stalked with an egg in the end like a stalked egg — somewhat as occurs in egg
masses of Scaphiopus hammondii.

At this writing (Dec, 1928) the fresh eggs of Aug. 16, 1922, now pre-
served in formalin surely look to be cream colored below and brown above.
Another accession of some fresh laid eggs in deeper water have the eggs ap-
pearing blackish. Furthermore, these eggs were preserved when mass was
still in the band form, the jelly unexpanded. Each band is from 4-6 eggs
wide. The width of some of these bands ranges from 5-8 mm. in breadth.

Dangers. Dr. Abbott emphasizes this point. "During this brief interval
these animals spawned, the eggs being attached to blades of grass and slender
twigs. These eggs hatched on the 2nd of July and a larger series were gath-
ered a week later.

'To return to the eggs. During the time that intervened from the laying
of the eggs until I gathered specimens of the tadpoles, there occurred four
moderately heavy showers, so that the water in the sink-hole at no time dis-
appeared, but was much below the level that it reached during the protracted
rainfall of June 26. Very much, therefore, of the spawn that was laid was
high and dry for from two to four days before hatching, and I suppose was
destroyed.

"On the evening of July 9 I found the water in the sink-hole confined to a
very few shallow pools of limited area, and in these pools were a few hundreds
of Scaphiopus tadpoles. In comparison with the abundance of eggs seen
June 26, and of young seen a week later, it is evident that a large portion of
the eggs were destroyed and a vast number of very young tadpoles were killed
by the soaking away of the water.

"I have never known any like disparity between the ages of frogs or com-
mon toads and the young in the tadpole state ; and it is at once very evident
that if the spade-foot habitually or usually deposit their eggs in temporary
pools, then we have an obvious reason for the positive rarity of the animal,
as apparently it iis the rule, rather than the exception, for the egg to be de-
stroyed or the young perish."

The eggs usually hatch before the water evaporates enough to do damage
except to the bunches which are near the shifting borders of these transient
pools. Few are infertile. A few masses so appeared and one mass we tried
to hatch so proved.

qS Frogs of the Okefinokee Swamp

HATCHING PERIOD

Andrew Nichols (1852, p. 115) writes: "In five days after I found the
spawn had become tadpoles, . . ." C. C. Abbott (1884, p. 1076) says:
"During this interval (June 26-28) these animals spawned, the eggs being
attached to blades of grass and slender twigs. The eggs hatched on the 2nd of
July. . . ." This makes 4 to 6 days for hatching. Sherwood (1896, p. 18)
gives the hatching period as "about a week."

On Aug. 4 and 5, 19 15 Dr. Overton (191 5, p. 53) records "a great con-
gress— enormous numbers of eggs of the spadefoot and Fowler's toads were
readily identified in the pools. On the 7th, the eggs were hatched" or 2 or
3 days after eggs were laid.

In this southern clime development is apparently even more rapid than in
the north. On Aug. 18, 1928, we have the field note that two lots "are hatched
early morning of August 18. In fact, some in pond were almost hatched
August 17 at noon or 1-1/4 days after laying. Certainly i 1/2-2 days after
egg deposition these have hatched. Water must have been 75° or 80° when
rain came and more when the sun came out. This hatched tadpole is from
dorsum very broad in head, has large gills, has body very narrow as compared
to head. The tail is dorso-ventrally very broad. The creature is mainly head
and tail and odd in appearance.

MATURE TADPOLE

Color description from life. (June 23, 27, 1922). General color bronzy.
Upper parts brownish drab, benzo brown, vinaceous drab. Upper parts with
many close set spots of apricot orange or vinaceous tawny which on the top of
upper tail crest become merged into a conspicuous bittersweet orange ridge
of color. Throat and gill region with little of the orange vinaceous spots of
upper parts. Gills can be seen through the skin. Spots on belly clear yellow
green in places.

Tail. Muscular part brownish drab, benzo brown, vinaceous drab. Crests
translucent. Upper crest with thin edge of dots and interlacing fines. This
arrangement goes around the tip. Rest of under crest translucent — no edging.
On the caudal half of lower edge of the muscular part the muscle segments
are more or less outlined by the fight color making emarginations between
the black dots. The same arrangement for the dorsal third of the muscular
part of the tail.

Iris and eye black, small; eye close together.

General appearance. Tadpole small (28.0 mm.) broad and large bodied
but not deep. Body has from dorsal aspect a LmM^MS-like appearance. Tail
short, tip blunt, rounded. Body unlike Bufo, broader nearer eye than vent.
Dorsal crest extends on to the body to a vertical just ahead of the developing
hind legs, is perceptibly nearer hind legs than spiracle. Spiracle sinistral,
far below lateral axis, as much or more ventral than lateral, opening more rec-
tangular or sfitlike than in our tadpoles in general. Spiracle about on the
level of the mouth. Eye on the lateral axis, very dorsal in position. The

Scaphiopus holhrookii 99

eyes close together, nearer the mid-dorsal axis than the lateral outline, in fact
nearer together than in Bufo. Anus median at the end of the edge of the
ventral cre^. Muciferous crypts in preserved s'pecimens at least indistinct.

Mouth parts: Teeth 6/6, 6/5, 5/6, 5/5. Whole labium (upper and lower)
with continuous row of papillae except on the upper labium where a pro-
nounced median interval is toothed. This median row is about equal to the
first lower labial row of either side. From upper labial interval of teeth down-
ward either side and across the lower labial border there is at least one row of
inner papillae. Opposite the ends of the horny beaks are several rows of
them filling in the space between upper and lower labia. The next series of
teeth (the second) below the upper toothed interval is usually not continu-
ous, rarely continuous. Usually one median end slightly overlaps the end of
its counter part of the other side. The third series has a short medium in-
terval one half the distance of the upper toothed interval or first series or
slightly more than a similar interval in the third lower labial series. Either
half of the upper third series is shghtly less in depth than the horny beak and
about equal to either half of the lower third series.

Measurements. Length of body (ia-12.2 mm.) in tail (12. 8-17. 6 mm.)
1.1-1.7, average 1.34. Width (6.0-7.4 mm.) of body in its own length 1.4-1.8,
average 1.64. Depth (4.8-6.8 mm.) of body i. 03-1. 50 in body width, aver-
age 1.23. Depth of body 1.73-2.5 in body length, average 2.04. Depth
(3.4-5.4 mm.) of tail in length of tail 2.5-4.0, average 3.14. Muscular part
(2.8-3.0 mm.) 1.45-2.4 in depth of tail, average 1.98. Spiracle i. 24-1. 66
nearer base of hind legs or vent region (4.4-6.0 mm.) than the tip of the snout
(7.0-8.0 mm.), average 1.41. Spiracle equidistant between eye 1.44-5.8 mm.
and base of hind legs or vent, 1.44-6.0 mm., average i.o. Eye 1.4-1.8 nearer
to tip of snout (3.0-3.6 mm.) than to spiracle (4.4-5.8 mm.), average 1.52.
Nostril 1.5-2.26 nearer eye (i. 0-1.8 mm.) than snout (2.0-3.0 mm.), average
1.9. Mouth (3.0-4.0 mm.) usually 1.56-2.5 larger than internasal space
(1.2-2.0 mm.), average 2.08. Mouth usually contained i. 0-1.66 larger
(average 1.3) than in interobital distance (2.0-3.2 mm.). Internasal space
contained in interorbital space i. 28-1. 83, average 1.56.

The dimensions of the largest tadpole are:

mm.

mm.

Total length

28.0

Spiracle to vent

6.0

Body length

12 .0

Spiracle to eye

5-2

Body depth

5-0

Eye to snout

3.6

Body width

6.4

Eye to nostril

1.8

Tail length

16.0

Nostril to snout

2 .4

Tail depth

4-4

Mouth

30

Tail, musculature of

30

Interorbital distance

30

Spiracle to snout

7-6

Internasal distance

2 .0

General remarks. After Nichols, Abbott is next to remark about the
tadpoles. He notes that "The tadpoles gathered July 9th, which were then
seven days old, were curious creatures. At this time the hind legs were well

loo Frogs of the Okefinokee Swamp

developed, although small, and did not interfere with the animal's natatorial
locomotion. The bodies of these young Scaphiopi were short, stout and oval,
and, when viewed in the water, deep velvety black; but when closely ex-
amined it was found that the two irregularly parallel yellow dorsal stripes,
that are so prominent a feature in the coloration of the adult male, were
plainly discernible.

"The movements of these tadpoles were not different from that of the
young frogs and toads in this stage of their existence. Those that I had in an
aquarium moved in companies as though following a leader, and occasionally
one would drop out of the ranks, come quickly to the surface, eject a bubble
of air, and dive again quickly to the bottom of the tank. Like all tadpoles
they had enormous appetites, and when fed with bits of raw meat quickly
attached their sucking mouths to the food offered, and did not remove it, I
think, while a particle of blood remained in the mass.

"A week later, July i6, the majority of these tadpoles had acquired their
front legs, and the tail had perceptibly diminished in size, but still was
used by them when moving through the water. At this time, however, the
movements of the animal are far less active than before or soon after, and for
a few days, if exposed to the attacks of any enemies, would suffer far more than
at any other period of their lives.

"A very curious feature in the growth of these animals is now to be
noticed. Of the specimens I had under examination, in an aquarium, about
five percent did not progress beyond the condition in which all were in July 9.
These 'retarded' tadpoles proved to be voracious cannibals. They seized
their more matured companions by their tails and legs, swallowing the meni-
ber and thus sustaining their own lives at the expense of their fellows. They
generally killed their victim in the course of twenty-four hours, and often in
less time, and then promptly seized another. So bloodthirsty were these
few 'retarded' tadpoles that I was compelled to protect the lives of the
little hoppers, their brethren, which now, in spite of stumps of tails, sat in
frog-like fashion on their haunches, and were in all respects miniatures of
the adult spadefoots that in April and June made night hideous with their
unearthly cries."

Pike (1886, pp. 219, 220) compares Spadefoot tadpoles to toad tadpoles.
His account pertinent to this topic is: "The first week I had them was warm
and sunny, and in about seven days the first tadpoles appeared, but a gloomy
cold spell following, the rest did not hatch out under twelve and even fifteen
days. At this stage they look black in the water, but are really black. In
about ten days the color broke out into blotches, with little white scattered
dots. Certain signs at this time led me to suspect I had found a treasure,
and the metamorphosis was watched with renewed interest. Having bred
toads, and my cabinet containing a series of these tadpoles of different ages, I
soon found a difference between them and my new acquisitions.

"The toad tadpoles are oval, of a dusky black, with a clear white fin on the
tail. The Spadefoots are brown and chubby, the tail narrow and blotched
all over. Then they swim differently; the latter with the body depressed,

Scaphiopus holbrookii loi

and they skim around the aquarium with the greatest rapidity. The tadpole
is of good size before the hind legs develop, and the back and abdomen
gleam with gold; the latter is dark gray, the former brown, with dark marks
on it showing the outline of the star of the adult starting from a broad dark
disk between the eyes.

"As soon as the hind legs are out, both body and tail diminish, and they
are as ravenous as other tadpoles, devouring both meat and fish greedily.
As soon as the thread-like front legs show, they must have cork or clips to
sit on, as their perfection is close at hand. I neglected this at first, and some
that were ready to leave the water August i8, actually atrophied till they
were barely an inch long from snout to hind toe, the smallest live reptiles I
ever saw, and died evidently from inability to remain in their watery home.
It was only by giving them resting places, and a way out into a dry house,
that I succeeded in bringing out strong young ones, and the first act of their
terrestrial life seems to be to provide a home by burrowing.

"When the front legs are well out in the toad, the whole under part
assumes a yellowish- white hue, the thighs are finely granulated, and on the
insides of the hands and feet the joints are thick and white. From between
the eyes and all over the back are the outlines of the future warts, increasing
in size daily. When the Spadefoot is at the same stage of growth the whole
underneath is dusky gray; feet and hands are slighter and smoother; the body
more drawn in behind the arms, and the tail is narrower."

In 1923 (p. 406) we held "Scaphiopus holbrookii has bronzy tadpoles,
translucent crests, rounded tail tip, oblique black bars on musculature; labial
teeth 4/5."

LARVAL PERIOD

Nichols (1852, pp. 115, 116) gives the first account of this phase: "In ex-
amining the water, however, we found it filled with spawn — and two females
were drawn from the bottom of the pool. . . . The old frogs were not seen
again. In five days after, I found the spawn had become tadpoles, of which
about a hundred I took home — kept them in glass globes, — fed them on fish
and flesh — scarcely one died. They remained longer and grew larger in the
tadpole state than did those left in their native pool, which later became
perfect animals in less than four weeks. On the other hand, those kept in
water, without any opportunity to crawl on land, or on any substitute for it,
such as floating chip, or some foothold firmer than water, were slower in their
development. It seems then, that the development and successive changes
of the organs, confirm to circumstances. So long as water is wholly their
residence, their caudal appendage is necessary and accordingly used, retain-
ing its proportionate size and strength, and the growth of the legs is in the
same proportion. If the water be gradually withdrawn, and mud, moist
earth and then dry, gradually substituted, they will much sooner undergo
the change from the embryonic to the infantile condition of existence. Thus,
at the end of four weeks, all the water in which the spawn was deposited had
evaporated; in some of the lowest spots of the basin a little mud of the con-
sistency of clay-mortar alone remaining. Here the young frogs were merrily

I02 Frogs of the Okefinokee Swamp

hopping about, enjoying life on dry land, while those in confinement in water,
were still increasing in size, and yet in an embryo condition. This very short
natural period of the tadpole state admirably fits them for such breeding
places as the one in which the subjects of this paper were found; a locality
where water can remain for a short time only, and this is the most wet seasons."

Sherwood (1898, p. 18) gives "the metamorphosis being complete in
about two to three weeks" or with his week of hatching 3-4 weeks from egg
laying. Ditmars (1905, p. 191) has it that "The metamorphosis is completed
within a few weeks from the time of hatching."

Overton (1914, p. 30) records for one group (19 13) that eggs laid Apr. 13
or 23 transformed the middle of June or two months later. In 191 5 eggs
which were laid Aug. 4 and 5, he found as transformed frogs at least by Sept.
4 or about 30 days from egg laying or 27 days of tadpole life.

Most of the records imply a very short larval period such as 30 days,
14-2 1 days, 2 1-28 days, 60 days, 30 days. Normally they need a month or less
before transformation. The eggs laid Aug. 16 and 17 were hatched August
17 and 18. Five days from egg laying these tadpoles were 14 mm. long, of
which the body was 7 mm. On August 26 we started northward in a Ford
and carried these tadpoles on the running board of the car. They neverthe-
less transformed Sept 2, or 17 days from egg deposition or strictly 15 days
of larval life.

TRANSFORMATION

Period. Nichols found them transforming in mid July. Apparently
Abbott's tadpoles under observation transformed in about 21-23 days or
July 23-25. Pike had them transformed in August. Overton records some in
September. In 1920 (p. 36) we held they transformed from June to August.

In 1922 we found in one place June 22 tadpoles, most of which were
mature, quite a few with two legs and one with four legs. On July 24, one
month later, there were none left in this pond. Transformation came from
June 2 2- July 24. Some eggs which were laid at Hilliard, Florida, August
16 and 17, 1922, transformed on the running board of the Ford Sept. 2.
The earliest transformation material we possess are some which Dr. Frank
Overton gave us in 191 5 bearing the data Aug.-Sept. 22, 1915, from Patchogue,
L. I. In view of the short larval period some trasnformations must begin
as early as mid April and some appear as late as October.

Size. In 1920 (p. 36) we give the "ranges of size from 0.3 to 0.5 of an inch
at the time of transformation (see Fig. 2 and PI. XXII, Fig. 11)." In the
figure 2 the transformation size is given as about 0.4 of an inch and the adult
as 2.7 inches. In the plate it is apparent the transformation spade-foot and
the common toad are the two smallest of the 16 species figured.

A series of 13 transformed spadefoots given to the writer by Dr. Frank
Overton, Patchogue, Long Island, and collected Sept. 22, 1915, rgfnge from
8.5-1 1 mm., mode 10 mm., average 10 mm. A transformed individual taken
June 22, 1922, at Camp Pinckney, Ga., measures 10 mm. A series of 17
taken at the same place June 27 give a range of 8.5-12 mm., an average of
10 mm., a mode of 11 mm.

Scaphiopus holhrookii 103

GROWTH

Colonel Nicholas Pike (1886, p. 218) writes that "When they leave the
water they are smaller than the common toad, but if the season is favorable
they grow rapidly in width more than length. I have some two years old
one-quarter of an inch in length, and nearly as wide as long."

We took in Okefinokee region transformation sizes from 8.5-12 mm. and
28 adults. The latter were all breeding males and females from 51-71 mm.
In the U. S. National Museum and our collections also intermediate sizes
between 51-12 mm. transformation size are scarce. We have a specimen
28 mm., one 44 mm., two 45 mm., and one 50 mm. Do the 28 mm. and 44 mm.
represent modes or two different groups? If so, we have four groups at least:
8.5-12 mm. at transformation; 28 mm., 44-50 mm., 51-72 mm. Doubtless
the last group needs to be divided into two groups. From comparison with
meager S. coiichii material where a 21 mm. specimen and 56 mm. were taken
at same place; and specimens 26, 27, 31, 31, 31, 31.5, 32, 36, 53, 56, 58, 59.5,
59.5, 60, 67 mm. were taken at another locality at the same time and from
deductions from 59 adults of S. holhrookii above 44 mm., the evidence pro-
visionally might be interpreted as: 8.5-12 mm., at transformation; 18-24 mm-
one year olds; 26-36 mm. two year olds; 38-50 mm. three year olds ; 51-61 mm.
four year olds; 62-72 five year olds. It could be interpreted as 8.5-12.0 mm.
at transformation; 21-28 one year olds; 31-40 mm. two year olds; 42-52 mm.
three year olds; 53-60 four year olds; 62-67 or 68 five year olds and 70-72
six year olds. At present we incline toward the former, knowing the material
is scant for careful conclusions.

FOOD

Holbrook alludes to its catching insects at the entrance to its hole. An-
drew Nichols (1852, p. 116) "also kept a few of the old frogs, three in number,
two females and a male in a barrel, a third part filled with moist peat muck,
containing some earthworms {Lumhricus terrestris L.) and other small crea-
tures. Occasionally I threw in a few garden snails {Limax agrestis L.),
small pieces of meat, fish or insects. Whether they ate of those provisions is
uncertain. Occasionally I found one or two of them out of the muck, in which
they usually buried themselves, reserving only a small breathing hole, opening
above their heads. ... I kept them until the last part of November,
when carelessly permitting the earth to become frozen, they were killed."

Pike (1886, pp. 217, 218) on Aug. 8, 1884 captured several of these animals
in their exodus from the water and makes some food notes on them. "We
carried our little treasures home in bags filled with grass, and so tired were we
after our hunt we left them imprisoned all night. Some of them objected to
being bagged, and made a faint squeaking noise when handled.

"Next morning I placed them in a large glass jar with earth, and in less
time than it takes to write the fact most were buried, all but their noses. I
fed them on flies and insects, and once I put in half a nest of young spiders.
As soon as their little bright eyes caught sight of the moving game, a most
animated scene took place. Every one was out jumping and capering about

I04 Frogs of the Okefinokee Swamp

till not a spider was left, when back they all hastened to hide till new victims
were provided. I turned many into the garden in the hope that I might see
some come out this spring. I was, however, disappointed, and it was not
until August 24 that by accident I discovered any were alive. On the night of
the 22nd, and all next day, heavy rain fell, which washed the poor little
fellows out of their homes and revealed their presence to me.

"Two days after the exodus we visited the same hillside, but with the
exception of two or three belated in the pond, not one was visible. We
hunted diligently for them, dug in all sorts of places, turned over heaps of
stones, but all in vain; yet I do not doubt there were hundreds buried all
around us had we only known where. It is evident they go far from their
watery home, for they were marching steadily on, the column spreading out
about fifty feet wide with none beyond it. Nor do I think they feed during
the exodus. I saw an immense number of very small crickets in amongst
them, but though I looked carefully I could not see any of the little Spade-
foots eating t,hem."

Fisher (1920, p. 77) had a captive of which he wrote: "While in cap-
tivity it was fed mainly upon earthworms. It also ate a grasshopper and a
cricket or two. It is remarkable how it would always close its eyes when
swallowing, and this seemed to be an important and necessary part of the act.
When open, the eyes bulge out prominently, but when closed they are drawn
back until they do not bulge at all. When the eyes are thus retracted, the
roof of the mouth is lowered and this doubtless helps to force food down the
throat in swallowing."

AUTUMNAL DISAPPEARANCE

Holbrook (1842, p. 1 1 1) says "I have met them even in very cold weather,
with snow on the ground." Col. Pike (1886, p. 215) writes of this species and
cold weather as follows:

"When the cold nights of fail begin, the Spadefoot leaves its summer home
and looks out for one more suitable for the winter season. It generally
chooses the warm southerly side of a hill, and excavates deeply for its new
quarters. It was only after many year's studying of this animal that I was
able to verify this fact.

"About four years ago I found one by accident in winter, over three feet
below the surface. On December 27, 1884, I was in Cypress Hills Cemetery
when a laborer who was digging a grave called my attention to a toad snugly
imbedded in the side of an opening he had just made. His spade had shghtly
grazed the body of the animal, which I saw at once was a Spadefoot. I asked
him not to disturb it till I had made a careful examination of the burrow. The
man had dug down nearly four feet, but the distance the creature had bur-
rowed was by exact measurement three feet two inches.

"The most careful search round the hibernaculum failed to discover any
outlet. It had left no trace of burrowing behind it, having evidently covered
up all tracks to its lair. The soil was closely packed about it, and the round
hole was perfectly smooth, just large enough to contain the body in the

Scaphiopus holhrookii 105

crouching posture I mention its assuming in confinement. I have since been
told that the Spadefoot has been found at a depth of six feet, but this I
cannot vouch for.

"When the weather is not too severe, even while the last snow is on the
ground, the Scaphiopus often makes its appearance, but then it only roams
aimlessly round, hiding under dead leaves and taking little food. The usual
time of awakening to renewed spring-life is the end of April or beginning of
May, and if the weather is unfavorable, not till June or even July."

Dr. G. Clyde Fisher (1920, pp. 76-78) says that "On August 14, 1918,
I captured a Spadefoot ... in the basement of my house in Douglaston,
Long Island, which is inside Greater New York. It was taken in a small
cavity in the damp earth, which it had probably made, in a break in the con-
crete on a level with the floor. It was in excellent condition, and apparently
it was fully grown. We had seen it hopping about at night several days pre-
vious to this, but we do not know how long it had been in the cellar or how it
got there.

"For several weeks we kept it in a large box of earth. At night it was gen-
erally out on top and active, but in the day time it was usually underneath
the soil. It would always dig in backwards, making good use of the horny
processes or spades on the hind feet. It was surprising how quickly it could
burrow out of sight.

"On the night of October 12 it sang a few times without being disturbed.
. . . It would usually sing or squawk when tickled on the throat or
breast.

"This specimen dug in on October 13 and stayed in so far as we could tell,
until dug out by us on October 21. It dug in on October 21, on which date
it ate an earthworm. On October 27, in order to make an experiment upon
its hibernation, it was placed out of doors, but unfortunately it escaped the
first night and was not seen again."

AFFINITIES

Holbrook's "General Remarks" (1842, p. m) are: "This animal is per-
haps somewhat allied to the Ceratophris of South America, which has teeth,
the posterior extremities short, and the hind feet furnished with a movable
unarmed tubercle. The Rana cultripes of Cuvier (Regne Animale Tom. II,
p. 105) would seem to be furnished with a process more nearly resembhng
that of our animal.

"The animals of the family Ranoidea deHght in the sun, and may at all
times be seen sitting half emersed in water, even when his days are the most
intense. They are all diurnal, or seek their food in the day time, with one
exception omy, the Scaphiopus solitarius, which passes its days in holes pre-
pared by itself, and feeds on such insects as may unwarily enter its dwelling."
Harlan (1835, p. 106) notes its likeness to Rana cultripes Cuvier. Dumeril
and Bibron, 1841, recognize its relation to Pelobates cultripes in their arrange-
ment. DeKay (1842, p. 66) holds that "With the teeth of a frog and parotid
glands of a toad, its natural place is between the.se two genera."

io6 Fj-ogs of the Okefinokee Swamp

LeConte (1855, pp. 429, 430) writes "This curious animal so much re-
sembhng a frog in its maxillary and palatine teeth, and a toad in its parotids,
the form of its body and its subterranean life, gives a fair example of some of
our systematic arrangements. It has been placed by M. Dumeril among the
frogs, of the genus Bufo. The fiat spur, as it has been called, at the root of
the first toe on the internal margin of metatarsus, is nothing more than a
much developed form of a scale or a disk occupying the same situation in
most animals of this family."

In 1855 Baird describes Scaphiopus couchii and in 1859 Scaphiopus ham-
mondii. In 1882 Boulenger places Scaphiopus holhrookii, Scaphiopus couchii
and Scaphiopus rectifrenis in the same portion of the key under "Derm of
head involved in cranial ossification" as opposed to "Derm of head free from
dermal ossification" in which group he places Scaphiopus hammondii. The
latter in 1866 Cope placed in Spea on the character Boulenger employs
above. In 1889 Cope still retained Spea for S. hammondii and S. multi-
plicata.

BIBLIOGRAPHY

1868 Abbott, C. C. Catalogue of the Vertebrate Animals of New Jersey. Appendix E.

in Geology of New Jersey by George H. Cook, p. 804.
1884 . Recent Studies of the Spade-foot Toad. The American Naturalist, Vol.

XVIII, No. II, Nov. 1884, pp. 1075-1080.

1904 Abbott, C. C. One Explanation of Reported Showers of Toads. Proc. Am. Phil.
Soc. Jan. -Dec. 1904, April 8, 1904, Vol. 43, pp. 163-164.

1868 Allen, J. A. Catalogue of the Reptiles and Batrachians found in the Vicinity of

Springfield, Mass., with notices of all the other species known to inhabit the State.

Proc. Bost. Soc. Nat. Hist., Dec. 2, 1868, Vol. XII, pp. 171-204.
1882 Boulenger, G. A. Catalogue of the Batrachia S. ecaudata. British Museum 1882.

PP- 433, 434-
1899 Boulenger, G. A. On the American Spade-foot {Scaphiopus solitariiis Holbrook)

Proc. Zool. Soc. London June 20, 1899, pp. 790-793.
1896 Brimley, C. S. Batrachia found at Raleigh, N. C. American Naturalist, Vol. 30,

June, 1896, p. 501.

1907 . A Key to the Species of Frogs and Toads Liable to Occur in North Carolina.

Journ. Elisha Mitchell Scientific Society, Vol. XXIII, Dec. 1907, p. 157.

1926 . Revised Key and List of the Amphibians and Reptiles of North Carolina.

Journ. Elisha Mitchell Sci. Soc, Vol. 42, Nos. i & 2, Oct. 1926, p. 80.
1889 Cope, E. D. The Batrachia of North America. Bull. No. 34, U. S. Nat. Mus., pp.

298-301.

1908 Davis, W. T. Notes on New Jersej' Amphibians and Reptiles. Proc. Staten Id.
Assoc. Arts and Sciences, Feb. -May 1908, Vol. II, Part II, pp. 48, 49.

1920 Deckert, R. T. Amphibian Notes from Dade Co., Florida. Copeia, Mar. 15, 1921,

No. 92, p. 22.
1922 . Notes on Dade County Salientia. Copeia, November 1922, No. 112, p. 88,

1905 Ditmars, R. L. The Batrachians of the Vicinity of New York City. The Amer.
Mus. Journal, Vol. V, pp. 190-191. Photographs by Hubert Lang.

1841 Dumeril, A. M. C. and Bibron, G. Erpetologie Ge'nerale, Vol. VIII, p. 472.

1920 Fisher, G. Clyde. A Spadefoot in Summer. Copeia, Aug. 18, 1920, No. 85, pp. 76-78.
1907 Fowler, H. W. The Amphibians and Reptiles of New Jersey. Rept. N. J. State

Museum, 1906, Trenton, pp. 92-97.

1918 . The Spade-foot Toad in Virginia. Copeia, March 19, 1918, No. 55, p. 44.

1858 Gunther, A. Catalogue of the Batrachia Salientia. London, 1858, p. 38.

1888 Hargitt, C. W. Recent Notes on Scaphiopus holbrookii. Amer. Naturalist, June

1888, No. 258, Vol. 22, pp. 535-537-
1835 Harlan, R. Medical and Physical Researches, Phila. 1835, pp. 105-106.
1902 Hay, W. P. A List of the Batrachians and Reptiles of the District of Columbia and

Vicinity. Proc. Biol. Soc. Wash., June 20, 1902, Vol. XV, p. 128.

1842 Holbrook, J. E. North American Herpetology, 2nd edition. Vol. IV, 1842, pp.
109-111.

1849 , In George White's Statistics of the State of Georgia. Savannah, Appen-
dix, p. 15.

Bufo quercicus 107

1856 Le Conte, John. Descriptive Catalofrue of the Ranina of the United States. Proe.
Acad. Nat. Sci. Phila., Dec. 185,^, Vol. VII, pp. 429, 430.

1852 Nichols, Andrew. Occurrence of Scaphiopus solitarius, in Essex County, with Some
Notices of its History, Habits, etc. Read June 17, 1843, Journ. Essex County Nat.
Hist. Soc, No. 3, 1852, pp. 113-117.

191 7 Nichols, J. T. Spade-foot Toad at Mastic, Long Island. Copeia, June 24, 1917,
No. 45, pp. 59, 60.

1927 Ortenburger, A. I. A List of the Reptiles and Amphibians from the Oklahoma Pan-
handle. Copeia, April-June, 1927, No. 163, p. 46.

1914 Overton, Frank. Long Island Fauna and Flora III. The Frogs and Toads. Mus.
Brooklyn Inst. Arts and Sciences, Science Bulletin Vol. II, No. 3, Nov. 3, 1914, pp.
28-30. '

19 1 5 . Annual Occurrence of Spade-foot Toads. Copeia, July 27, 19 15, No. 20,

P- 17-
1915 . Late-Breeding Spade-foot Toads, etc. Copeia, November 18, 1915, No.

24, pp. 52, 53-
1886 Pike, Nicholas. Notes on the Hermit Spadefoot — . Bull. Am. Mus. Nat. Hist.,

Vol. I, 1881-1886. Bull. No. 7, Art. XIV, July 1886, pp. 213-220.
1865 Putnam, F. W. Some Statements Concerning the Frogs and Toads found about

Cambridge, Mass. Proc. Bost. Soc. Nat. Hist., Vol. 2, (1862, 1863) 1865, pp. 229-230.
1898 Sherwood, Wm. L. The Frogs and Toads Found in the Vicinity of New York City.

Proc. Linn. Soc. of New York, 1897, 1898, No. 10. Read Apr. 13, 1897, pp. 17, 18.
1879 Smith, F. S. The Spade-foot Toad in New Haven, Conn. American Naturalist,

Oct. 1879, Vol. XIII, No. 10, pp. 651, 652.

1923 Van Hyning, T. A Collecting Note on Florida Batrachians. Copeia, May 20, 1923,
No. 118, p. 68.

1920 Wright, A. H. Frogs: Their Natural History and LTtilization. U. S. Bur. Fisheries,

Appendix VI, Doc. 888, 1920, p. 22, 27.
1923a . The tadpoles of the frogs of Okefinokee Swamp, Georgia. The Anatomical

Record, Jan. 20, 1923, Vol. 24, No. 6, p. 406.
1923b . The Salientia of the Okefinokee Swamp. Copeia, Feb. i, 1923, No. 115,

P- 34-
1926 . The Vertebrate Life of Okefinokee Swamp in Relation to the Atlantic

Coastal Plain. Ecology VII, No. i, January 1926, p. 83.

1924 Wright, A. H. and A. A. Wright. A Key to the Eggs of the Salientia East of the
Mississippi River. The American Naturahst, Vol. LVIII, July-August 1924, p. 379.

Bufo quercicus Holbrook

(PI. I, Fig. 5; V, Fig. 7; VIII, Fig. 2; X, Fig. 13; XV, Fig. 12; XVII; XX; XXI; XXII,
Fig. 7, Text Fig. I, 4)

COMMON NAMES
Oak Toad. Dwarf Toad. Oak Frog.

RANGE

Check list. Type locality: Charleston, South Carolina and Smithville,
North CaroHna (Loding says this is now Southport).

Range. North Carohna to Alabama and Florida — Stejneger and Bar-
bour 1923, p. 26.

Supplementary records. Since 1923 it has been recorded in Mississippi and
Louisiana. In 1922 H. P. Loding (p. 17) records these "Alabama records:
Mobile and Baldwin Counties," In 1923 Viosca (p. 37) records it in South-
eastern Louisiana. In 1924 (p. 59) G. S. Myers takes it in Wilmington, N. C.
The same year Schmidt (1924, p. 68) has it from Mt. Pleasant, S. C. The
same gentleman took it at Natural Well, Magnolia, N. C, on July 10, 191 5.
In 1926 (p. 80) C. S. Brimley gives it as an inhabitant of Craven, Carteret,
Duplin, Edgecombe, Lenoir and New Hanover Counties, N. C.

io8 Frogs oj the Okefinokee Swamp

In igo8 (p. 19) Brimley held "Dwarf Toad (Bufo quercicus) "to be in the
Lower Austral Zone. In 1926 (p. 82) Wright held ''Bufo quercicus N. C. —
Fla. — Ala." to be one of the eleven species of frogs of the Okefinokee region
which might be held to be in the Sabalian Zone or region. The same year
1926 Percy Viosca Jr., in species of "Group I, Species Common to the Atlantic
and East Gulf Coastal Plains" writes "Excepting where governed by strictly
local ecological conditions, we can consider it axiomatic that species found
on the Atlantic Coastal Plain occur at least as far west as the Florida parishes
of Louisiana, an area lying in the extreme Southeastern corner of the East
Gulf Coastal Plain. My own observations have already extended westward
the known range of a number of species, the following of which can be in-
cluded in Group I: Bufo quercicus, Hyla gratiosa, Abastor erythrogrammus,
Leimadophis fiavilatus and Tantilla coronata. Those which have not pre-
viously been recorded as far west as Louisiana, I have taken in St. Tammany
or Washington Parishes."

Local Okefinokee records. In 191 2 we wrote of it as follows: Twenty-four
specimens were taken in 191 2, sixteen May 28, 191 2, on the trip into the
swamp, three from May 3- June 2, two June 6, and three June 24. They
were secured on Billy's Island, Honey Island and at Mixon's Ferry. Their
occurrence in Okefinokee comes well within their range. There is no definite
place given for the Georgia records which LeConte (Cope '89, p. 292) made
in 1855. This species seems most common in Florida, where Brimley ('11,
p. 11) added four stations to Cope's three records for the State and Miss
Dickerson ('07, p. 105) has it from Ozona, Fla. It reaches its northern limit
in North Carolina.

In 192 1 we took it on every island visited (Billy's, Floyd's, Honey, Black
Jack, Jones, Middle, Chesser and others); in the country surrounding the
swamp, such as The Pocket, from Hopkins to Waycross, from Folkston to
Waycross, along the Suwanee River to Fargo and along the St. Mary's River
from Moniac to Camp Pinckney (in 1922 from Camp Pinckney to St. Mary's
at its mouth). It is a very abundant and universally distributed species.

GENERAL APPEARANCE

Holbrook (1842, Vol. V, p. 13), its describer, gives its characters as follows:
"Head short; snout pointed; superciliary arches slightly elevated; body short,
very fiat, rounded at the flanks, above dusky, with a yellowish vertebral hne,
on each side of which are black blotches ; abdomen silvery-gray ; throat dusky ;
groins tinged with yellow." DeKay (1842, Vol. Ill, p. 68) speaks of this
extra-limited species as "very small." Cope (1889, p. 292) speaks of it as
"the smallest known species of the genus Bufo."

C. S. Brimley (Dec. 1907, p. 157) has it as "Size small, length of head and
body one inch. Skin very rough. Bony ridges turning inward almost at
right angles just back of the eyes." In 1926 (p. 80) he characterizes it as
follows: "Size small, length of adult only i 1/4 inches, a white line down
middle of the back, skin very rough especially on legs." Cope (i88g, p. 261)
thought "One smaller metatarsal tubercle; superciliary crests incurved pos-

Bufo quercicus 109

teriorly so as to be transverse; a supratympanic crest; parotoid gland descend-
ing on sides to inferior part of the tympanum," — the diagnostic characters of
Bufo quercicus. Miss Dickerson (1906, p. 45) emphasizes the "Cranial crests,
inconspicuous; skin very tubercular, spinous on legs and arms; size small
(i 1/4 inches)." Both LeConte and Loennberg and others have found that
the superciliary crests can be straight at times.

The identifications of the oak toads in the past have emphasized their
smallness, their scarcity in most collections, the paucity of good accounts of
them, and other factors. For example, it is not surprising to find a young
Scaphiopus holhrookii identified as Bufo quercicus or the young of other Bufo
species thus termed. We have seen as many as ten different accessions in
collections of earlier years dubbed "Juvenile," "Young," "Half-grown"
when in almost every instance they were full grown males and females ex-
ternally apparent as such. The collectors did not distinguish them as oak
toads nor were they used to seeing the species.

COLORATION OF SPIRIT SPECIMENS (19 1 2)

Coloration. — Upper parts greyish through light to dark brown or almost
blackish. From the tip of the snout to the vent runs a sharply defined
white or yellowish white line and even in the darkest-colored individuals. In
the series of comparable small southern toads this vertebral color is absent
or if present, is irregular and primarily along the middle third of the back.
There are 5 or 6 pairs of black spots along this line. Usually from the nostrils
of either side there extends backward a black or brownish bar which makes a
right angle turn to the front of the eyelid; the second pair begins on the
posterior part of the eyelid and extend rather obliquely to the vertebral line
and along it inside of the supraorbital crest to its end. These two pairs of
black or brown spots produce on the head a cross of intervening white color
one bar being the median longitudical vertebral line and the other a trans-
verse line from the middle of one eyelid to the other eyelid. Sometimes just
inside the anterior end of the parotoid and somewhat away from the vertebral
line is a pin point of a spot comparable to the same one in B. terrestris and
B.fowleri. On a line with the posterior ends of the parotoid are two other
large prominent spots which may be longitudinal or oblique in relation to the
median line. Halfway between the posterior ends of the parotoids and the
vent is another pair of spots and just ahead of the vent a sixth pair. This
sixth pair may be absent at times as are the 3rd pair. In some of the fighter
colored specimens from the posterior end of the parotoid there leads backward
along either side a prominent black band bordered above by an almost
equally wide white band. Also from the front part of the parotoid a dark bar
leads downward to the arm insertion. All of these spots and the whole dorsal
coloration pattern is very suggestive of the detailed color description of Bufo
fowleri (Dickerson, '01, p. 94) and of some of our B. terrestris already de-
scribed, but if these specimens be compared with B. terrestris representatives
of the same size, the latter have the spots more irregular, usually more or
less obscured by reddish brown tipped warts, smaller spots back of the

no Frogs of the Okefinokee Swamp

posterior ends of the parotoids. The fourth pair of spots of B. quercicus have
their counterparts in B. terrestris in two pairs of spots, the anterior pair
pin-head spots which usually meet and fuse across the middle line.

The underparts are grey or greyish-v/hite spotted or unspotted. Most of
our specimens fall in the former group. Possibly Florida specimens (like
our St. Petersburg material) may be unspotted but all of our specimens but
two are more or less spotted and we have from Natural Wells, Magnolia,
N. C, a specimen collected by Mr. Karl P. Schmidt, which is spotted and
even darker beneath than our Okefinokee forms. In some the spotting is
very faint on the breast and pectoral region. In others it occupies much of
the upper belly and one has all the underparts dark except the very wrinkled
oval patch on the buttocks which is pure white or yellowish-white. Usually
when dark enters the ventral coloration the throat is darkest. Most of these
specimens have the palm and sole dark but not all of them, and our young
southern toads have these parts white in the main except in a few which are
dark colored.

STRUCTURAL CHARACTERS (WRITTEN IN 191 2)

Structural characters. — In the structural characters we have our best
distinctions between the oak toads and immature southern toads. The
throat pouch or vocal sac of the male is distinctive but sexual characters are
tantalizing key distinctions unless given for both sexes. In the cranial
crests these differ from Bufo terrestris at the same size. In Bufo terrestris
there are no crests or the supraorbital crest is very faintly outlined. In B.
terrestris these ridges decidedly bend inwards at the posterior ends and the
interval between these two converging ends is contained 2.5-3.0 times in the
length of the crest to the nostril while in B. quercicus the ends are divergent
or very slightly convergent and the interval is contained in the crest to the
nostril 1.5-2.0 times. Whenever the crests show in these 20-35 mm. southern
toads the posterior end of the continuous supraorbital crest is invariably
beyond the inner posterior-dorsal angle of the upper eyelid and opposite or
behind the anterior end of the parotoid while in B. quercicus adults 20-30 mm.
it ends ahead of the anterior end of the parotoid and frequently stops at the
above mentioned angle. Behind this angle it frequently extends as a series
of interrupted raised warts. In none of the southern toads are the posterior
supraorbital crest ends connected across the middle line by a ridge, or series
of warts, but is always open while in the oak toads there is always a transverse
series of raised warts which give the cranial hollow a parapet behind as well
as on either side. In very warty specimens the other crests loose the dis-
tinctiveness and only this parapet of three sides shows. In southern toads
20-35 mm. there are no postocular and supratympanic crests at all. In oak
toads the postocular ridge is never continuous with the supraorbital crest and
the gap is bridged by a series of warts. The supratjTnpanic crest from the
postocular to parotoid is very prominent and very broad in B. quercicus while
in small B. terrestris (20-35 mm.) it is absent and in 50-70 mm. adults it is
always a smooth edged horny ridge not with numerous warty tubercles on it.

Plate XX

Oak toad {Bufo quercicus)

1 . Ilex mijriifoUo — Bufo quercicus pond, about four miles E. X. E. of Chesser Id.,

Ga. July 15, 1922.

2. Male croaking, Billy Id., Ga. July 3, 192 1. Flashlight.

3. Male croaking, Billy Id. Ga. May 26, 192 1. Flashlight.

4. 5. Male croaking, Billy Id., Ga. June 4, 1921. Flashlights.

6. Bvfo quercicus on sand, Honey Id., Ga. July 3, 1921.

7. Egg string, attached to leaf blade. Coat-bet pond, Chesser Id., Ga. July 17, 1921.

Bufo quercicus in

The parotoids of the oak toad are finely spinose while those of a small southern
toad are no more than tuberculate at the most ; the parotoids of the former are
relatively shorter their width i. 25-1. 5, usually 1.5 times in the parotoid's
length, while in the latter the parotoids are more elongate, 1.8-2.0, usually
2.0 in their length. The posterior ends of the parotoids of B. quercicus are
far more divergent than those of B. terrestris, i.e., set more obliqufely to the
vertebral line. The parotoids of B. quercicus in many send a downward por-
tion to the lower level of the tympanum while in these young B. terrestris, as
in adults, the lower edge of the parotoid, is little if any below the upper border
of the tympanum. The tympanum is in contrast with the postorbital ridge.
Upper parts including upper eyelids rugose, on the fore-hmbs and sometimes
on the tarsus, spinose; symphyseal tubercle as in young southern toads;
underparts granulate in young and females, rugose often in the larger males;
first finger shorter than second and about equal to the 4th ; outer metacarpal
tubercle quite large, inner tubercle small; subarticular tubercles often in a
double series, the tubercle at the base of each finger transverse or double in
nature and as wide as finger tips; the other sole tubercles and subarticular
ones are circular. Outer metatarsal tubercle small; inner tubercle large and
brown-tipped; plantar tubercles relatively smaller than the palmar tubercles.
In all except six of the 24 specimens the tongue is small, narrow, elongate
and elliptical with the posterior third or half free. In one it is as broad as
long; in four quite broad and much shorter and in one specimen the front half
is broad and the free half narrow.

MEASUREMENTS

(1912-1914)

The measurements of 8 adults are as follows : In length they are 2 1-30 mm.,
average 25.3 mm., the head including the tympanum is 7-9 mm., average 8
mm., usually 3 sometimes 3.5 times in the length of the body, or head meas-
ured to rictus oris is 4-4.6 in the length, or the head measured to posterior
end of the supraorbital crest is 3.3-4.3 in the length; width of the head 7-10.5
mm., average 9 mm., usually i.i times greater than the length of the head,
equal to the femur and tibia; snout 2.75-4 mm., average 3.3 mm., less than
the eye which is 3.5-4 mm., average 3.5 mm. while in small B. terrestris
of the same size usually the two are equal; tympanum indistinct, 1.25-2 mm.,
average 1.65 mm., much less than the interorbital distance or eyehd; inter-
orbital distance 2.25-3.5 mm., average 3 mm., usually greater than the
upper eyelid which is 2-3 mm., average 2.5 mm.; femur from 6.5-10.75 mm.,
average 9.35 mm., slightly more than tibia which is 7.5-10 mm., average
8.8 mm., femur slightly more than foot without tarsus; tarsus 4.5-6.75
mm., average 5.75 mm.; rest of foot 6-9.75 mm., average 8.3 mm., equal
to or shghtly less than the tibia; anterior limb from axilla 9-16 mm., average
12.2 mm.; posterior limb from groin 18-29 mm., average 22 mm. or forward
on body to posterior part of the posterior part of the parotoid or to the arm
insertion.

In all these measurements we compared these B. quercicus specimens with
about 40 B. terrestris of the same size and season of capture, and must confess

112 Frogs of the Okefinokee Swamp

we could find no constant relative measurement of the above categories to set
one apart from the other. It seems that usually the breadth of these oak
toads in the length is 1.5-2.0 while with almost all of the small southern
toads of the same size the ratio is 2.0 or more, but this is not always absolute.

(Recent Material)

Head to angle of mouth 1.6 (20 mm.cf ) — 1.6 (28 mm. 9 ) — 1.5 (28 mm.cf )
in width of head; head to rear of tympanum 1.14 — 1.07 — 1.24 in width of
head; head to angle of mouth 4.0 — 4.66 — 4.0 in length of body; head to rear
of tympanum 2.85—3.1—3.3 in length of body; snout .71— -75— -75 in first
finger; snout i.o — i.o — .75 in fourth finger; snout .43^-37 — -37 in first toe;
eye 1.4 — 1.33 — 1.14 in snout; eye .4 — .66 — .57 in tympanum; eye 1.0 — 1.0 —
.87 in first finger; tympanum 7.5 — 4.0 — 4.25 in intertympanic width; tym-
panum 3.5 — 2.0 — 2.0 in snout; internasal width 1.33 — 1.0 — 10. in upper
eyelid width; interorbital width 1.0 — .83 — .71 in upper eyehd width; inter-
orbital width .75 — .83 — .71 in internasal width; interorbital width 3.75 — 2.66
■ — 2.4 in intertympanic width;

Forelimb: Forelimb 2.0 — 2.0 — 1.93 in length of body; forelimb 1.9 — 1.85
^1.8 in hind limb; first finger 2.2 — 1.66 — 1.66 in third finger; second finger
1.83 — 1.4 — 1.66 in third finger; second finger .83 — .86 — 1.0 in first finger;
third finger .45 — .70 — .70 in second toe; fourth finger 1.6 — 1.25 — 1.66 in
third finger; fourth finger .43 — .375 — .5 in first toe; internasal width 1.66
— 1.2 — 1.2 in first finger; internasal width 2.0 — 1.4 — 1.2 in second finger;
internasal width 3.66 — 2.0 — 2.0 in third finger; internasal width 2.33 — 1.6
— 12. in fourth finger.

Hindlimh: length .95 — .92 — .92 in hind limb; tibia 2.87 — 2.66 — 2.8 in
length; tibia 2.7 — 2.5 — 2.6 in hind limb; tibia 1.42 — 1.33 — 1.45 in forelimb;
tibia .91 — .95—1.0 in hind foot; first toe 1.66 — 2.33 — 2.33 in second toe; first
toe 3.0 — 4.0 — 3.33 in third toe; first toe 4.0 — 6.0 — 6.0 in fourth toe; first toe
2.66 — 3.33 — 3.0 in fifth toe; second toe 1.8 — 1.7 — 1.43 in third toe; second toe
2.4 — 2.55 — 2.1 in fourth toe; second toe 1.6 — 1.4 — 1.3 in fifth toe; third toe
1.33 — 1.5 — 1.4 in fourth toe; third toe .9 — .83 — .9 in fifth toe; fourth toe
1.08 — I.I — 1.33 in hind foot; fourth toe 1.16 — 1.17 — 1.33 in tibia; fourth toe
1.66 — 1.56 — 1.9 in forelimb; fifth toe 1.5 — 1.8 — 1.7 in fourth toe; internasal
width 1.0 — .60 — .60 in first toe ; internasal width 1.66 — 1.4 — 1.4 in second toe;
internasal width 3.0 — 2.4 — 2.0 in third toe; internasal width 4.0 — 3.6 — 3.0
in fourth toe; internasal width 2.66 — 2.0 — 1.8 in fifth toe.

HABITAT

Holbrook (1842, p. 14) found that "This beautiful little species of toad is
mostly found about sandy places that are covered with a small species of oak
which springs up so abundantly where pine forests have been destroyed;
whence it is commonly enough called Oak Frog, which specific name I have
preferred". At this time he limited it to South Carolina and North Carolina
but in 1849 (Appendix, p. 15) he gives "Bufo quercicus — Oak Frog" as one of
the Batrachia of Georgia.

LeConte (1855, p. 430) held it to be "Very common in Georgia in wet
places, under logs and pieces of wood".

Bujo quercicus 113

Loennberg (1895, P- 5.^^) in 1892 and 1893 found it "very abundant on the
dry sand hills about Oakland, Orange County, as well as in the 'flatwood'
about Kissimee, Osceola County and in Hillsboro County".

Brimley (1909, p. 133) of Lake ElHs region had "About a half dozen taken
in drier situations in May 1908 (also taken by Sherman on Shackleford's
Banks, near Beaufort, N. C. in June 1901.)" The same author (1910, p. u)
"received (it) from Hastings, Orlando, Green Cove Springs and St. Peters-
burg, all in Florida. Evidently common at the first and last two places".

Ten miles south of Jacksonville, Florida, in 191 1 and 191 2 R. L. Deckert
makes the observation that "Bufo quercicus Holbrook (is) common, in com-
panies in temporary water, or scattered on cultivated ground".

In Alabama Loding (1922, p. 17) writes "In Mobile County this little toad
is rather common in high sandy locations with Scrub Oak and Palmetto under-
growth".

In Louisiana in 1923 (p. 37) Viosca places this species in his "Division 4,
Longleaf Pine Flats" of "The Uplands". This Division he characterizes as
follows: "Abruptly bordering the pine hills on the south, lies this strip of fiat
lands characterized by a beautiful stand of longleaf pine interspersed with
loblolly. The streams here, because of the lower elevation, are normally slug-
gish and often spread out over low areas forming swampy situations. These
are characterized by swamp magnoHa in the shallower areas and tupelo in the
more permanent swamps. — Typically the country is north Floridian, rather
than Louisianian, and several Southeastern species, not found elsewhere i'n
our State, have congenial habitats here. Bufo quercicus, Bufo terrestris and
Hyla gratiosa are significant examples".

In 191 2 we wrote of their habitat somewhat as follows: "This little toad
was found on the outskirts of the swamp in the cut-over lands where few pines
remain and where the cover was solely low heaths. Here at noon when we
arrived these creatures were hopping about little mindful of the sun. On
Honey Island we took them on the sandy soil where the pines were abundant
and on Billy's Island they occurred on the unshaded cleared cultivated fields
of the Lee's. They seemed to have no particular preference for shade or ex-
posure and apparently were quite generally distributed. The few specimens
in our collection are not due to their scarcity so much as to our neglect. One
might think them quite terrestrial from their spinose exterior but not only on
the drier and higher outskirts did we find them but also on the islands and
near their swampy edges. In fact, we even recorded some on the little islets
of Honey Island Prairie."

After our 1921 and 1922 studies we think of this form as an essentially
abundant pine barrens species though occurring in hammocks and elsewhere.

FIRST APPEARANCE

In 191 2 we found them when we entered the swamp the last week in May.
In 192 1 we record them as early as April 25 when we entered the region and in
1922 we arrived June 13-14 when they were out in full force. Some of the
U. S. National Museum material was taken in April, such as:

1 1 4 Frogs of the Okefinokee Swamp

U.S.N.M., Nos. 29056-64, Apr. 3, 1901, adults, L. Kissimee, Fla., E. A.
Mearns.

U.S.N.M., Nos. 49580-81, Apr. 16, 1912, Autaugaville, Ala.

U.S.N.M., No. 46147, Apr. 20, 1892, Mobile Bay, Ala., R. J. Thompson.

U.S.N.M., No. 46090, Apr. 23, 1892, Bay St. Louis, Miss., Mr. V. Bailey.

One record is for March, 191 2, made by N. R. Wood at Auburndale, Fla.,
(U.S.N.M. Nos. 48771-76).

GENERAL HABITS

Variation in color. Holbrook (1842, pp. 13, 14) characterizes this species
as follows:

"The head above is dusky, with a yellowish central longitudinal Hne; the
superciliary ridges are grey, with a white mark in the centre; on each side of
this longitudinal line is an oblong spot, extending from it to include most of
the posterior part of the orbit of the eye; a small part only of the orbit in front
of this is light coloured; the upper jaw is light brown.

"The back of the animal is dusky-brown, with a vertebral line of pale
yellow, marked with a few scattered small warts of reddish-brown colour; on
each side of this line are irregular black blotches, with here and there a sHght
tinge of reddish-brown; the back is covered with innumerable warts and
granulations, of variable size and colour, generally black, but the smaller ones
of dusky-red ; on each flank, and extending from the axilla downwards towards
the posterior extremity, is an oblong black blotch, bounded with white both
above and below.

"The throat is dusky; the abdomen is silvery-grey, yellowish at the groins,
and with a pale tinge of yellow around vent. The anterior extremities, as well
as the posterior, are dusky-brown above, marked with black transverse bars
or spots; their inferior surface is coloured like the abdomen, except the fingers
and toes, which are reddish-brown".

For coloration of our 191 2 material see topic Coloration in Spirits (1912),
also the topic Mating — Colour — Male and Female. They may be any shade
of grey, brown or reddish-brown or even be almost black. Loennberg
noticed that some of his material from Hillsboro County were redder. Ap-
parently his other material was grey or light brown. LeConte speaks of it as
"dusky with a few irregular black spots, many of the warts, particularly those
on the sides whitish or reddish ; a very distinct line of one of these colours runs
from the point of the nose along the vertebral column to the vent. Beneath
the body is granulate, white, more or less varied, particularly on the fore part
with black". Cope with his mistakes on this species we do not quote. Miss
Dickerson (1906, p. 104) gives a good colour description and accompanying
plate of the species. This species if it wishes can be as variable as Bufo
terrestris or Bufo americanus.

General habits. Holbrook (1842, p. 14) held "it spends most of its time
in concealment under fallen leaves, or partially buried in the sand, from which
it is washed out by heavy rains".

Bufo quercicus 115

Loennberg (1895, P- 33^) says "This is a very active little animal con-
sidering the fact that it is a toad. It is seen in all kinds of places and at all
times of the day, even in the brightest sunshine, but especially after rain".

VOICE

During the breeding season according to Holbrook (1842, Vol. V, p. 14)
"the male (has) a slight chirp, not unlike some kinds of insects". To Deckert
(1914b, p. 2) "its cry resembles that of a small chick, very loud and shrill, and
may be heard at any time of day or night, sometimes from absolutely dry and
dusky fields and roadsides, where these tiny toads hop about in the glaring
sunshine, living on the smallest of insects. The full chorus, which is ear
splitting, is, however, heard only at night, after heavy rains".

]Miss Dickerson (1906, pp. 19, 21, 105, 106) speaks of Bufo quercicus thus:
"Vocal bladders inflated from the middle of the throat are to be found in
Bufo cognatus, Bufo compactilis and Bufo quercicus''. ^'Bufo quercicus would
seem to be expressing most active distress in its tones like those of a lost
chicken". "They are difficult to see, but give notice of high-pitched sounds.
The individual call is like that of a young chicken in distress, but considerably
louder. The male alone gives the call, and while producing it seems to have in
his mouth a transparent bladdder about the size of a man's thumb. The fact
is that this toad has a large vocal bladder that can be extended from the
mid-line to the lower throat region. This structure relates it to Bufo com-
pactilis and Bufo cognatus of the Southwest. When taken in the hand, the
Oak Toad gives a rather musical chirping sound, like that of a young bird".

The first day we recorded its calling in 192 1 comes May 16. Then one of
the lads brought in an oak toad he had taken on the edge of a cypress pond.
A little later Mr. Harper "went off in the piney woods. He came back and
reported one of the biggest queries he has yet encountered. He said he didn't
know whether the caller was a bird, or beast, or whether it might be young
bobwhites, queer, brownheaded nuthatches or some other bird? A rain came
up and not until he was just ready to leave did he solve the note. It was that
of Bufo quercicus. It seemed so unfroghke or untoadlike a call to him".

His journal note is as follows: "A baffling, peeping in several parts of the
piney woods. Some strange Hyla, bobwhite or something althogether differ-
ent? Nearly desperate over these peepings here and there in the pinej' woods,
always stopping when I came close. — Scarcely know whether to look in trees
or on the ground. Finally one peeped in a fairly open, burnt over space, and
by hearing it from several directions at close range, I hit upon the spot pretty
closely. Then getting on my knees I had scarcely gone a yard, when a little
black toad, with a thin, golden-yellow stripe down its back, made a Httle
movement among the grass and saw palmetto. Bufo quercicus! And mean-
while other peepings continue at intervals in the other parts of the piney
woods; pheep, pheep, pheep, pheep, pheep, pheep, pheep, pheep, pheep, pheep,
pheep, — a plaintive (rather shrill) high-pitched piping.

"When I 'froze' for several minutes, it quit 'freezing" on its part, and made
a few leisurely forward movements, crawhng with one leg at a time, lizard-

ii6 Frogs of the Okefinokee Swamp

like, with body off the ground. It made hops of a couple of inches or so only
when I poked a straw at it. Meanwhile little loose fold of skin in the middle
of its throat vibrates. Once in a while it seems to nab an insect.

"A cloudy, lowering, humid day, following a day and night of rain. —
Finally it made off of its own volition, — hops of two inches or so, though it
still mainly crawled. It was interested in ants running around near it, turning
its body to watch them and finally thrusting out its tongue to lick one up — ".

The afternoon of this day, according to my own journal, we "went out to
Crosby Pond. — Where the road crossed a swale we heard a queer note and
approached it from several angles. We found its author near the base of a
stump in amongst saw palmetto, gallberry (Ilex glabra) and Gaylussacia.
The oak toad male was calling before we approached. We got within ten feet.
He piped only lowly. After we had worked him around for a photo for some
time, he suddenly to our surprise backed into a hole at the base of a saw
palmetto. The hole was 3/4 of an inch in diameter and not deep. Once when
one of us made a pass at a fly the toad ducked farther into its hole. Its note is
surely very birdlike. One will hear three or four calls like a piping chicken.
Sometimes the notes are repeated three or four times. Then process re-
peated after a very short interval. There may be three or four groups of
calling for one individual. There were several animals around. Once the
note was likened to that of a swallowtailed kite. Truly the most unfroglike
note I ever heard. It is very high pitched and some times sounds like some
animal in distress. There are several calling in the piney woods. One caUing
from a tangle of chokeberries (Aronia), Osmunda cinnamonea, Bamboo brier
(Smilax) and sweet bays. Couldn't find it. Are the toads moving pond-
ward? Later in the evening we heard none". The next day the author
visited this toad's burrow. When we ran a straw into it only a queer spider
ran out. The toad had left.

In 1912 and in 192 1 some of the residents both normally and almost in-
variably accurate assured me that the blacksnake had a whistle and that this
note of the oak toad was the call in question. They readily acquiesced in our
determination of it as that of Bufo quercicus.

On June 4, 1921, while the author was studying eggs Mr. Harper made the
following field notes on peeping and intervals in Bufo quercicus. In the after-
noon are the following: "Number of peeps during a calling period are about
25, 16, 13, 32, 37, 35, 32, 24 and 27 (for one observation). Only a few brief
seconds between periods. One interval of 3 seconds; then records of about
35 peeps in 20 seconds, 16 calls in 8 seconds; 18 calls in 10 seconds; 30 calls in
15 seconds; 32 calls in 16 seconds; 29 calls in 14 seconds". "In the evening
they were slower — perhaps exhausted with 4 to 5 hours of continual calling.
10 calls in 26 seconds; 22 in 16 seconds; 20 in 13 seconds; 22 in 17 seconds".

In inflation of the throat they are like Bufo compactilis, Bufo cognatus.
The lower throat is the principal part involved in the process. The lower
part of the throat is thrown out into an elliptical bag or sausagelike
balloon. One can tell when a toad is going to thrill after a rest. The body
will inflate to a large size and then the most ludricous sac projects out in the

Bufo quercicus 117

throat region. The tip of the sac when not really inflated comes close to the
tip of the chin. Otherwise it appears as a little loose vibrating sac 1/2 inch
out from the lower part of the throat. When deflated the body inflates. When
body is compressed or deflated the sac inflates.

Its call may carry 1/8 of a mile or more. When this species is in chorus it
is one of the most deafening ear deadeners I have encountered when in their
midst. We often employed the method of varying the closure of our ears to
pick out various notes and cut out other dominant notes. For example, Mr.
Harper's note of a chorus 8:30 — 10:00 P. M., July 16 illustrates it "Bufo
quercicus, abundant calling. All {Hyla gratiosa, Chorophilus, Hylafemoralis,
Bufo quercicus) combined, enough to cave in our ear drums. By half closing
ears we shut out the sound of Hylafemoralis almost entirely, and hear others,
better than ever especially Chorophilus, Bufo quercicus, Acris, — ".

The journal voice records for 192 1 and 1922 are as follows:

1921

May 16. First heard to be recognized. Ventriloquial. Many heard.

May 17. Heard one or two more.

May 18. Heard sone B. quercicus.

May 21. After afternoon rain temperature 71° heard several oak toads.

May 22. Several heard noon-day sun. Only one heard near Oak Toad swale.

May 23. At 5:30 and 7:30 p.m. heard one near camp.

May 24. Hottest day yet, 94° at 3:00 p.m. Heard one, then later several.

May 26. Are Bufo quercicus bound for the ponds? Have taken and heard several. This
afternoon when hottest an occasional Bufo quercicus spoke.

May 27. One calling 4:30 p.m.

May 28. One calling sundown.

June I. Hear oak toads particularly near trestles. In cypress ponds hear occasional oak
toads.

June 3. None calling this night.

June 4. Rained 2 inches or more. Here at congress. Heard plenty amongst saw palmetto.
Lots croaking in the ponds.

June 5. Several calling during and after the shower. Few this evening.

June 6. No B. quercicus calling.

June 17. One heard.

June 20. One heard. From time to time we heard oak toads.

June 21. One heard.

June 22. Heard at 9:30 p.m. one oak toad.

June 23. In white quarters heard several croaking. Thunderstorm after evening began.
Bufo quercicus calling.

June 25. Several heard 9:30 a.m. Hear none near camp.

June 26. One heard 6:30 p.m. Temperature 77°. Showery.

June 28. Rain in evening. Commonly calling.

June 29. Heard before and after thunderstorm.

June 30. Afternoon. Heard as I approached Billy's Island 6-6:30 oak toads in consider-
able numbers. Few calling near camp.

July I . Few calling near camp.

July 3. Calling during and after rain. Numerous everywhere.

July 4. Evening at 9:00 p.m. 75° sultry. No B. quercicus calhng.

July 5. One recorded in the evening and one in mid-afternoon.

July 6. One or two in afternoon after rain. Several calling in temporary pools.

July 15. One heard.

July 16. Abundantly calling at night.

July 19. 10:00 A.M., one heard.

July 23. Few Bufo quercicus calling.

July 24. One 12:30 p.m. (rain). Later calling commonly.

July 26. Quite a few calling at night.

July 29. Commonly heard in rain from Honey Island Prairie to The Pocket.

July 30. No end of oak toads calling. Rain in late afternoon.

July 3 1 . Heard abundantly along the railroad in a good many places from Fargo to Moniac.
Heard also practically every night along the St. Mary's — F. Harper.

Aug. 17. Bufo quercicus several heard at Camp Pinckney.

i8

Frogs of the Okefinokee Swamp

1922

June 13. Calling at 11:45 bright sun. Air about 94°.

June 15. Several calling on mainland.

June 19. Heard a few in woods after the rain of today.

June 21. One calling at 10:00 p.m.

June 22. Heard around camp a few calling.

June 23. Heard at R.R. ponds one mile south of Trader's Hill. Afternoon.

June 26. One calling. Cloudy weather. Cutover pine barrens. Several about Anna's
Pond. Several on way to Folkston and back to camp.

June 27. One heard at 10:00 a.m. in cut pine barrens.

July 2. 8:30 P.M. — midnight. Heard a few. Some rain.

July 3. Last night to midnight or later in many places a deafening roar of Bufo querdcus.
It began before dusk.

July II. On the island heard oak toads everywhere at 5:00 p.m. after the rain. Tempera-
ture 70°.

July 12. Several calling this forenoon.

July 18. One or two heard at Murray Bay.

July 21. One or two heard in mid-forenoon. Showery.

July 25. Heard a few in mid-forenoon. Bright sun. Hear plenty calling west edge of
woods at 9:00 in evening. Air 72°. Hard rain until 7:00 p.m.

July 26. Immense chorus on mainland and on Chesser Island. Tonight at midnight 75°
big din.

Aug. I. One calling mid-afternoon. Many heard at night.

Aug. 2. One or two heard at about 11:00 a.m.

Aug. 4. Several in pine barrens 8:00 p.m.

Aug. 5. Several heard tonight.

Aug. 8. No end of oak toads heard along old Okefinokee road 8:00-1
They are everywhere.

Aug. 9. Heard from 5:00 p.m. onwards at mainland and about camp. Common.

Aug. 10. Commonly heard near Petty Pond.

Aug. II. Tonight 8:00-12:00 heard quite a few oak toads from Folkston to Chesser Island.

Aug. 13. St. Mary's Ga., — Chesser Island. Strong Chorus 7:00-10:30 p.m.

Aug. 15. One or two late afternoon.

Aug. 16. Commonly calling in spade-foot ponds at night.

Aug. 17. On oak ridges near and around our camp never saw more oak toads. Some calling.

Aug. 20. One heard at daybreak. Temperature 72°.

: 00 P.M. after rain.

In 192 1 on the day of the record of calhng the air minima of nearby
stations (5 in all) were from 6o°-7o° in May, 6i°-78° in June and 67°-73° in
July or an average of 68°. Our camp records usually taken when we arose in
the morning range from 64°-83° or average 73°.

In 192 1 and 1922 when great congresses came, the following rain and
temperature records for five nearby stations are as follows. The rain con-
ditions at camp are in the last column.

1921

Degrees

Degrees

May 16

S-47

inches

81-83

66-67

Heavy rain

" 21

3

13

"

79-80

66-67

Continual rain

June 3

4

04

nches

82-87

68-70

Sprinkles

4

70

83-88

64-71

Downpour

5

82

82-90

69-70

Shower

" 30

3

37

81-89

66-67

Thunderstorm

July 3

4

25

86-91

70-72

Rain all P. M.

" 16

I

16

85-87

70-72

Shower

" 23

2

97

" 24

I

30

87-90

67-70

Rain

" 29

4

66

73-84

70-72

Downpour

" 30

I

74

85-95

71-72

July

Aug.

Bufo quercicus

115

1922

2

5.31 inches

90-96

68-72

Heavy rain P.M.

II

1. 16 "

87-92

66-70

" " "

25

■73 "

93-96

73-74

26

T. "

94

73-74

Rain P.M.

8

3.64 "

85-93

73-74

9

•55 "

81-93

71-73

Showers

10

1.24 "

79-91

68-73

Sprinkles

II

.87 "

85-91

65-69

13

2.79 "

87-91

68-73

Showers

16

2.0s "

85-91

70-72

Rain most P.M.

" 17 4-53 " 82-91 69-72 Cloudy

In 192 1 in every instance rain came in nearby stations and at camp (one
exception) when congresses came. The stations average .14-1.10 inches per
station, the maxima range from 79°-95°, the minima 66°- 72°. In 1922 rain
came in every instance in nearby stations from i to i .06 inches per station or
always at camp (4 exceptions). The air maxima were 8i°-96°, air minima

66°-74°.

With these subterranean animals, like the spade-foots, warm rains at least
0.5-1.0 inches or very humid conditions seem needful to start it calMng
vigorously. It may start at minima of 6o°-67° but the minima of congresses
are usually 66°-74°. Humidity is, however, the important factor.

MATING

Coloration {From life, June 8, 1921 ) . Dorsal stripe white, pale orange yellow
or maize yellow or sulphur yellow or cream buff. The dorsal spots are black.
First pair between eyes usually not connected as in Bufo terrestris; spot in
front of parotoid round one wart, not elliptic with two or more warts; spot
between parotoids not connected, not a half moon made by connection; spots
ahead of hump separate as in B. terrestris; spots ahead of vent prominent
enough in oak toad but just two pin points. Upper parts with some gull gray
or pearl gray or pale olive gray. Stripe from lower part of tympanum almost
to groin in the above grays; also patch back of angle of mouth below tympan-
um also patch above arm insertion, in front of femur and back part of upper
eyelid. All these lighter dorsal areas with burnt sienna colored tubercles
which are especially prominent along either side of dorsal stripe from
hump backward and on oblique lateral stripe, on posterior part of eyelid. The
tubercles on black areas look black but many are really also burnt sienna.
Parotoid with fine and thickly studded burnt sienna tubercles. Tubercles on
palmar and solar surfaces, posterior surface of thighs (partially), groin (a lit-
tle), pectoral region (a few tubercles) vinaceous rufous. Hay's russet or mars
orange. Ventral parts smoke gray or grayish white or pale olive buff or even
sometimes a cream color. Each tubercle stands out. On venter proper they
are close together and black between, little apparent on the sides, tubercles
wider apart and the intervening black more apparent. Same for underside of
the limbs.

I20 Frogs of the Okefinokee Swamp

Iris cream color, rim around pupil and eye napthalene yellow; rest largely
black with some cartridge buff or ivory yellow.

Male with lower throat between angles of mouth dark olive gray, more or
less puffed out, pleated or wrinkled. All of throat darker than rest of ventral
parts but lower half darker than mental part. Female with throat same as rest
of venter. Slight dark spot in anterior posterior region.

Structural differences. Neither Holbrook nor LeConte indicated the sexual
differences. Miss Dickerson was the first to indicate that the throat of the
male is dusky. Deckert (1914b, No. 9, p. 2) says "The throat of the male is
black, and when the pouch is distended, forms a small elongate bladder,
slanting upward, and about 3/4 inch long and 1/4 inch in diameter".

In 191 2 we made these notes on the males and females. "These four females
are the lightest colored of the twenty-four, the upper parts being grayish-
brown and the spots being most distinct. In one of the spent females the
underparts are pure white and the other gravid female has only a few faint
spot's but the males are the darker specimens. In all of these a gular fold or
flap extends acrosss the lower throat connecting the posterior ends of each
ramus of the lower jaw. In the larger males when the vocal sac is not dis-
tended it lies behind this gular line, and looks like a triangular apron on the
pectoral region the apex pointing backward and the base being on the gular
line. On the females this gular barely shows. In the males we could discover
no excrescences on the first two fingers. This breeding evidence is one of the
strongest arguments to prove that this little animal is not a spinose young B.
terrestris or americanus as Boulenger thought, and is corroborative of Hol-
brook's third reason for its establishment (Holbrook, 1842, V, p. 15)."

A study of our 191 2, 192 1 and 1922 specimens and such other Bufo
quercicus material which has come under our observation resolve our data in
the following categories:

1. The range of our males from Okefinokee is from 19-30 mm., of males from
widespread parts of its range (U. S. N. material) 20-29 mm,

2. The range of our females from Okefinokee is from 20.5-32 mm.; of other
females from U. S. Mus. collections (26 lots) elsewhere 22-28 mm.

3. The grand average of all females measured is 25 mm., mode, 26 mm. The 69
females measured were as follows: i at 20.5 mm.; 3 at 21 mm.; 7, 22 mm.; 9,
23 mm.; 4, 24 mm.; 12, 25 mm.; 13, 26 mm.; 8, 27 mm.; 6, 28 mm.; 2, 29 mm.;
2, 30 mm.; i, 31 mm.; i, 32 mm.

4. The grand average of all males measured is 23.6 mm., mode 25 mm. The 89
males measured were as follows: 5 at 19 mm.; 4, 20 mm.; 3, 21 mm.; 11, 22
mm.; 7, 23 mm.; 11, 24 mm.; 22, 25 mm.; 12, 26 mm.; 5, 27 mm.; 5, 28 mm.;
I, 29 mm.; and i, 30 mm.

5. The measurements of 11 actual pairs are somewhat as follows:

June 4, 1921 22cf X 259 ; 26cf X 30.59 ;

July 3, 1921 25cf X 26.59 ; 23cf X 24.59 ; 24cf X 269 ■.2sd' X 269 .

July 3, 1922 25.50^ X269;26cr X 289 24cf X319.

July 26, 1922 26.5cf X 279 ;

July 27, 1922 25.5cf X 25.59 .
In no instance was female smaller; but the average is no more than i 1/2 mm.
greater in the females, the greatest range of difference 7 mm., the least o mm.

Biifo quercicus 121

6. Several notes on males Or possible males might be in place:

(a) 15.5 mm. specimen has i transverse fold from mouth angle to other
mouth angle. Is it to be a male? At this size these small oak toads
are little beauties: stripe down back very prominent from mouth
margin over snout to vent. Tubercles carmine tipped and at times
making a margin along light vertebral line.

(6) 16 mm. has 3 transverse edges across lower throat. Is it going to be
a cf ? cf not functional at this size.

(c) Of 4 specimens 14, 15, 16, 17 mm. we remark: two small ones darker
in color and will probably develop into cf s. No signs now. The
larger two are lighter, may develop into cf s.

(d) One 18 mm. specimen black throat, two transverse folds, one of them
connecting angles of mouth; probably going to be cf .

(e) One 19 mm. cf , throat in general not black, the evertible sac not
extening over pectoral region as lapette or apron. It is outlined in
front by two transverse ridges and behind in pectoral region by two
transverse ridges.

(/) One 19 mm.c?', throat region with 5 or 6 transverse folds but no apron.

(g) One 19 mm. cf , lower throat with good apron on breast.

{h) One 19.5 mm. cf , dark throat and narrow dark triangle down middle

of pectoral region.
(i) One 20 mm. cf, throat not very dark but has throat lapette.
(j) One 20.5 mm. cf , quite a throat apron. Do you suppose some queer

oak toad noises are young males calling?
(k) One 22 mm. Lapette extends forward beyond chin.
(l) One 24 mm. Dark throat apron. No black excrescences revealed on

thumb and second finger in alcoholics,
(m) One 24.5 mm. Possibly first and second fingers in life may seem to

be a little more horny but it is not very well revealed in preserved

material,
(n) One 26 mm. Apron on lower throat but throat light not dark while

its mate, a female, has dark throat and heavily spotted breast.
(0) One 28 mm. Dark throat. Lapette covering 3 or more folds under-
neath. One or two ridges on front of pectoral region hid by apron.

No excrescences revealed. Forearm and possibly brachium in males

thickened and relatively larger in this male. Probably a general rule.
(p) One 30 mm. Beautiful. Long lapette with secondary one. Top of

head reminds me in some ways of Bufo hemiophrys.

7. In a similar way a few notes on some females are :

(a) 18 mm. specimen 9 ? but not a male.

(6) 18 mm. 2 transverse ridges on lower throat. Is it a 9 ?

(c) 21.5 mm. ripe, dark throat.

(d) 22 mm. with dark triangle on pectoral region.

(e) 25 mm. pectoral region with prominent black spots.

(/) 25 mm. Very dark spotted throat and breast. (To distinguish from cf
must look at throat vesicle. Other female throats like rest of under
parts. Tympanum warty or may be free of warts. Sometimes tympanum
in female so covered with warts tympanum is not revealed.) 9 ripe.
Surely a warty beast.

(g) 25.5 mm. spent, 13 mm. wide.

(h) 25.5 mm. ripe. Stripe more prominent in preserved 9 s, probably
because they are lighter,

(i) 26 mm. Dusky throat and breast.

(j) 26.5 mm. Throat slightly spotted with darker.

(k) 26.5 dark, ripe. Width 19 mm. Triangle on pectoral region.

122 Frogs of the Okefinokee Swamp

(l) 26.5 mm. Ripe. Underparts generally light throat slightly darker.

Width 15 mm.
(??i) 27 mm. Throat dark extended on pectoral region and front belly as a

triangle, the base of the triangle on the throat.
(n) 27.5 mm. Ripe. Stripe down back almost bluish-white in alcohol.

Under parts unspotted.
(0) 28 mm. Solid black throat, heavily black spotted breast. Spent,
(p) 30 mm. Ripe. Width 18 mm. Underparts uniformly light.
(q) 30-S mm. Dark throat and vestee. Spent. 14.5 mm. wide,
(r) 31 mm. Throat dusky. Middle of pectoral region with several big
black spots. Width 20 mm.
8. In general it seems that males are revealed externally as such at 19 mm. or
higher and females at 20.5 mm. or higher, that this is two years from trans-
formation, i.e., the oak toad goes through two winters after transformation
before breeding.

Duration, day or night. In 192 1 almost the instant, April 25, we entered
the swamp the boys brought us a cf oak toad. But it was not until May 16-2 1
we began to record the females. On the latter date we noted that ''in the
afternoon after the rain we started for turpentine still. Heard and saw Bufo
quercicus. Picked up two, one a gravid female". Later "went to old and new
hogshole. Heard a few males in the water. They all stopped except one which
I could not find. On May 25 we took 2 males and 2 females."

On May 26 we made the following journal notes: "In pipewort, sedge and
grass places at 10:00 a. m. found a female Bufo quercicus. Hear males in the
woods. Is B. quercicus actually going to the ponds soon? B. quercicus calls
are more lively and insistent. Have taken four or five this morning. Females
are about more since last night's thunderstorm. Are females bound for any
particular place on this piney island? They are moving somewhere. Does it
mean the onset of breeding? In burnt-over area it seemed as if more were
present. Possibly they are easier to find in this area. Found 3 cf s and 39 s.
Males not in holes. Am getting so I can locate them easier. Returned to
camp to find that the boys had taken 6 Bufo quercicus from the hammock
west of camp. One male mated with another, made a terrarium for the oak
toads. Now all 12 or 14 are out of sight in sand except two. They back into
ground".

On June i we were still puzzling about them per the following: "An oak
toad beside a log; very hght in color; can squeeze into small space and freeze.
It was near a ditch. Found two oak toads by a log near the trestle but still on
dry ground. They are a pair but not mated. Will they go to water separately
or mate on the way without rain or mate with first rain? Now have a plenty
of them, 15 or 16".

Only three days later, June 4, many of our queries were answered. About
two inches of rain dropped and the island seemed teeming with oak toads (see
Ovulation for account) . They bred almost everywhere. All about the cleared
fields, in piney woods, in hammocks, and numerous other places we found oak
toads that day. In one pond heard plenty of male oak toads amongst saw
palmetto. We saw lots of Bufo quercicus croaking on floats, one on a pine cone
in the water, others on logs, sticks, etc." "In one case a little cf oak toad

Bujo quercicus 123

faced a large southern toad {Bufo terrestris). Both were croaking and their
inflated sacs would almost touch. A picture indeed. AVe found several pairs
on the south side of this pond. They were in water 4-6 inches deep and easy
of capture. They do not separate. Could have taken more in this little
transient pond where there are no end of B. quercicus^'. On June 5 near our
old camp of 19 1 2 in a pond where vigorous matings were held on June 4
found a dead female, no doubt killed from rigors of mating or from ex-
haustion of ovulation or from both causes.

On June 28 at night we found Bufo quercicus calling vigorously. Francis
Harper relates the following amusing incident: "A Bufo quercicus, on a float-
ing chip, had to leave one foreleg suspended in air when 'hollering', the dis-
tention of body preventing the toad from reaching a foothold with it. Another
seen on floating chips."

On July 3 "the species was abroad in great numbers. Every transient
shallow pool filled by the rain had them calling. Took 3 or 4 pairs and 30 or
40 males in short order".

Amplexation {Normal, cross, ahnormal). In all the amplexations of
field mated pairs and also matings in captivity the typical axillary form ob-
tained. Sometimes ovulation may come without attendant male. For
example, "July 26, 1922, at midnight heard no end of Bufo quercicus in pond
near Starling Branch crossing. Captured two pairs in an overflowed grassy
area. One pair when we arrived at camp were broken. Put them together in
fish can but they did not mate. Probably female laid her complement without
mating. The other pair laid by morning of July 27".

In several instances we noted among our captives that a male might mate
with another male. On July 5, 1921 "in my enclosure found three in tier
arrangement with a female below, embraced by a male in axillary fashion and
this male embraced by another male in typical axillary amplexation.

OVULATION

Habitat. "In the breeding season, they seek out stagnant pools, where

they deposit their spawn, (Holbrook, 1842, Vol. V, p. 14). In 1920 (p. 30)

we state that "the egg string is laid in warm, shallow ponds, . . . ". In 192 1 we
found them in any kind of impermanent pools after a heavy rain. They might
be on the swampy edge of the island or in the highest normal dry parts of the
island, might be in railroad ditches, excavations, shallow pools or ponds,
cleared fields, corn patch puddles, depressions in roads, or any basin which
will hold water for a time. At times, however, one is impressed with the ob-
servation that possibly they do discriminate permanent places. For example,
on June i, 192 1, we observe: "Hear oak toads particularly near trestles.
Do they know there is more permanent water in such places?" Many of them
breed in shallow cypress ponds where development is assured.

One journal note (1922) will suffice to indicate their widespread distri-
bution and choice of breeding. "July 3. Last night in many places a deafen-
ing roar of Bufo quercicus. Especially at a pond 1/2 mile west of Trader's
Hill, Ga., — a shallow grassy area. There were four Florida tree frogs ( Hyla

124 Frogs of the Okefinokee Swamp

gratiosa) calling, some lo southern tree hogs (Hyla squirella), some cricket-
frogs (Acris gryllus) and no end of pine woods tree frogs {Hyla femoralis).
Never heard such a din. Hard on the ear. Brought in 12-15 males of oak
toads {Bujo quercicus). Also brought in three pairs of oak toads". The above
species are frequent associates of the oak toad in breeding time and place.

Period. In 19 12 we discussed the ovulation period from scanty evidence.
It follows: Of all the 24 adults only four are females, three of which are spent
and one of the four, the largest specimen (30 mm. long), has the ovaries full of
eggs about to be laid. This specimen was taken June i, just one day previous
to the circumstantial record in the field and these four records give us little
idea of the breeding range. Subsequent to June 2 we found no more of these
small egg-strings nor ripe females and these facts force us to believe June i
about the end of the egg-laying period. Five of the eight specimens were
taken from June 1-7 and breeding impulses may explain their greater abun-
dance during the first of June and not later.

In 1920 (p. 30) we held that "the oak toad breeds in May and June, most
of the egg laying doubtless occurring before June 15". In 1923 (b, p. 34) we
write from June ist to loth, three species begin (ovulation) Bufo quercicus . . . ".
In 1924 (b, p. 379) we give as the "Season June 4 to August 21".

In 192 1 on June 2 we noted that the "oak toads from Hopkins, Ga., have
not laid as yet. The females have ripe eggs in them".

On June 4 we found our first eggs. It "rained" two inches or more. When
we came out from dinner the island was afloat and everywhere oak toads were
calling. Oak toads out where apparently no permanent ponds are but never-
theless in these ponds calling and mating. Found a pond where they were
calling and laying '.

The next congress came on July 3, 1921. Our terse note is "70° at 7:35
a.m. Trip by pond lever car to Honey Island, Honey Island Prairie and al-
most to Black Jack Island. On return to Honey Island began to pour, but a
warm rain. Found oak toads in a shallow brushy basin. Immense choruses.
Caught several. In fact, caught about 50. We took three pairs of oak toads.
In an overflow area with Woodwardia (fern) the predominant plant, water 2-4
inches deep. This chorus at 3 : 30-4: 00 p.m. Same condition all the way home".

In scanning our journals, work sheets and notes we find notations as to
ovulation on the basis of different factors as follows: May and June, May i-
June 15, May 25- July 30, June i-August 21, June 4 to August 21, June 4-
August 27, June 4- August 17, May 25- , June 4- August 8, June 4- July
17. These dates give the growth of our accumulating data but they also
reveal that our knowlege of this abundant species is all to scanty. Doubtless
if one were to choose the extremes above May i-August 2 7 he could not be far
astray, although the author would prefer May 15-August 15 or June i-
August 15 for the general period and bulk of ovulation.

If one occasionally finds mature tadpoles in mid June or unspent females
(August 26, 1922) in late August then he realizes the possibility of May i or
earlier as beginning of oviposition and September possibly as the end of
straggling ovulation.

Bufo querciciis 125

Since the above paragraphs were written we have examined very carefully
our female series and the U. S. National Museum material with the following
results: Spent May 23, May 25, June i, June 4 (3 9 ), June 5, June 6, June
15, June 23, July 3, 4 9 , July 4, 2 9 , July 13-17, 2 9 , July 15- , July

27,2 9 and August 8; unspent-May 29, June i, June 2, June 15, June 16, 5
9 , June 28, June 30, July 3, July 12, 2 9 , July 15- , July 17, July 26,

2 9 . The range of ovulation on this basis might then be from May 23-July
26, a possibility, August 8. The May 23 date is from St. Petersburg, Fla., and
one unspent record in peninsular Florida extends at least to July 17, — these
stations south of the Okefinokee Swamp.

Temperature and humidity. In 192 1 and 1922 when positive ovulation was
known in seven instances, 3 in 1921 and 4 in 1922 our temperatures at camp
range from 7o°-82° and in every instance when rains from showers to drenches
obtained. The minima of surrounding weather stations are from 64°-74°.
In ovulation often the heavier part of the rain may have come the day before.
The following night the immense chorus or congress may ensue. In June and
July when a rain starts about noon and continues through the afternoon quite
certainly that evening if minima be not below 60-64 the oak toads appear en
masse. The minima of the 7 ovulation records 7o°-82° in camp are somewhat
higher than the minima recorded and derived for congresses, i.e., 64°-74°
(see voice) but are exactly the same as temperatures above (64°-74°) derived
from nearby weather stations for the seven dates.

Egg-laying process. Holbrook (1842, Vol. V, p. 15) writes 'T have seen the
male and female together, and have seen the female deposit her spawn, even
when confined in a glass vessel". He does not, however, describe the process.

On June 4, 192 1, the first congress we ever witnessed we did not see all the
process. Some of the pairs laid eggs in a bunch reminding one of Chinese fire-
crackers (see egg description). "The oak toads began laying at 10:00 p.m.
The female would emit some eggs. The male would draw up his legs as in B.
terrestris and B. americanus and emit milt with two or three exertions. Then
he would rest in drawn-up fashion for i or 2 minutes or sometimes less. Then
eggs would drop down. At first one pair laid eggs only in bars of 2 or 3 or 4 or
5 or 6. Seemed so strange. Female then would lay 14-30 eggs at one time or
4-6 bars. Then once after egg-laying she dragged along and all the bars
trailed behind showing they had a connection, a hyaline jelly connecting them.
There is no outer envelope as in B. americanus and apparently eggs come out
in bars of 4-6, then area between becomes stretched. At times several bars
or free ends appear from the vent of the female and not just two ends, i.e.,
one from each oviduct".

For a month after first eggs in laboratory we did not see eggs in the field.
We wondered if they were heavier than water, sunk to the bottom and were so
small we could not find them on the bottom.

On July 3 and 1 7 we began to get our clues. On July 3 we found oak toads
common in the furrows between rows of newly planted sweet potatoes. In
the flooded furrows were no end of toads. Saw an oak toad pair swim to the

126 Frogs of the Okefinokee Swamp

bottom of a furrow in its deepest part and they deposited eggs. They moved
on. They don't leave all eggs in one place. Saw 7 or 8 oak toad pairs. All
amplexate axillary fashion".

EGGS

Attachment, egg mass. Holbrook, though he saw the female lay her spawn
does not describe the eggs, their form and characters. In 19 12 we made the
following provisional identification:

On June 2, 19 12, in a shallow pool in the hollow between two growing corn
rows we found the small egg files of some kind of toad. At the time we had no
preservative or cans to take care of these and upon return the next day the
pool had dried up. Our one prominent impression was the smallness of the
egg string. The eggs were in one row like those of our northern B. americanus
and rested on the muddy bottom of this shallow impermanent pool, but the
egg-tube was much smaller in diameter and length. The egg string was not
carefully examined at the time and we cannot compare it with the eggs of the
common toads. We are, however, firmly convinced that these files were those
of the oak toad. Deckert (1914b, No. 9, p. 2) speaks of "The spawn . . . laid in
tiny strings, . . . ". In 1920 (p. 30) we speak of "The egg string or file is a small
edition of the southern toad's egg string, the former being much smaller in
diameter than the latter". Deckert and we each should have used bars or
short strings, for it better conveys the form of the egg mass. In 1923 (p. 406)
Wright and Wright speak of "Eggs in files. Bufo quercicus . . .'\ In 1923
(b, p. 34) Wright holds "Bufo quercicus lays its short bars of eggs on the
bottom of ponds".

In 192 1 our first field notes on eggs came June 4. They are: "Found a
pond where male oak toads were calhng. Bufo quercicus eggs. Many parts of
an egg complement will be two, three, four, five or six eggs in a string or bar
independent of rest. One pair, at first laid nothing else but these bars. Often
times s or 6 will radiate from one center. These are all in same plane. Some-
times from one center bars extend in all planes, e.g., one of 13 bars; another of
30 or 40 or 50 bars. The jelly envelope closely fits the egg and the margin
becomes slightly indented between each egg. The eggs are black and white;
last night by flashlight the eggs looked creamy but fear we were wrong on this
observation. Egg more brown or brownish-black on upper pole. Apparently
the bars of eggs is comparable to the inner tube of B. americanus eggs and the
tube of jelly becomes pinched off or drawn out between each four or five eggs.
Sometimes eggs separating to bars, sometimes not. An egg bar will be from
2-3 mm. to 6 or 7 mm. in length. Vitellus i.o mm.; 9.5 mm., 9.0 mm.; .8 mm. ;
i.o mm. Whole bar 5.8 mm., leaves 1.15 mm. envelopes between on both ends
or about 2 mm. between each egg. A partition line between each egg".

On July 3 we did not get an entire clue to field deposition but on July 1 7
on Chesser Island "In water 3-6 inches deep with dip net discovered one or two
strings of 4-6 eggs of Bufo quercicus. Became accustomed to finding the 4-6
egg bars on the bottom without dip net. The single bar or two of them seems a
regular deposition form in the field".

Plate XXI

Oak toad (Bufo quercicus)

1 . Habitat near Siren Run, Billy Id., Ga. May 27, 192 1.

2. Toad looking out from entrance of hole in ground, Billy Id., Ga. May 16, 192 1.

3. Toad in daytime retreat i^ inches deep and Ys inch in diameter, beneath a slab

of wood. Trail Ridge, Folkston, Ga. June 16, 1922.

4. Female and male, Chesser Id., Ga. July 4, 1922. Ventral aspect, x i.o.

5. Cluster of egg bars laid in camp, Billy Id., Ga. July 4, 192 1.

6. Egg bars, stained, Chesser Id., Ga. July 4, 1922. x 1.3.

7. 8. 9. Tadpoles, Chesser Id., Ga. August 10, 1922. x 1.0.

[o. Transformed toad, Chesser Id., Ga. July 23, 1922. Dorsal aspect, x i.o.

Bufo quercicus 127

Egg description. In 191 2 we made these observations on the species. A
count of the eggs of the left ovary reveals 305 eggs and if there be the same in
the right one the complement would be 610 eggs. Each of these ovarian eggs
is .8-1.0 mm. in diameter, the average .9 mm. while in the common toad the
egg is 1. 0-1.4 mm., average i.i mm., mode 1.2 mm.

In 1920 (p. 30) we held that "The female oak toad may deposit 500 to 600
eggs, while a common toad produces 4,000 or more. The eggs of the oak toad
are slightly smaller than those of the common toad". In 1924 (pp. 576, 377,
379) Wright and Wright give a diagram of a bar of 5 Bufo quercicus eggs quite
unhke in egg string any Bufo east of the Mississippi. Of all Bufo eggs seen in
the U. S. A. they remind us most of Bufo compactilis only its eggs are in long
strings. Our characterization of 1924 (p. 379) is "Eggs submerged. Eggs in
files. Files short (4 to 10 mm. in length) ; 4 to 8 eggs in short bead-hke chain
or bar or many such files radiating from one focus; vitellus 0.8-1.0 mm.; tube
diameter 1.2-1.4 mm. Egg complement, 610, 766".

In 192 1 we first positively identified this species (See egg mass, attach-
ment, for 192 1 notes). Two days after the July congress we observe that we
made this note. "Noticed that the eggs of Bufo quercicus are decidedly
brown and cream or yellowish".

In 1922 the following three notes might have pertinence: On "June 28 we
found a female in a 'chufa' field at 9:00 p.m. Later made a count of the ripe
eggs in her left ovary. There were 383 or if the same for the right, 766 for
both ovaries, its entire complement". On "July 4 the pairs laid between 12
midnight and 7 :oo a.m. The strings come out several eggs long but the jelly
every 5 or 6 eggs seems to pull out. There is little jelly around eggs. There
is httle jelly apparent if stained with red ink or potassium permanganate
(camp stains for writing and poison ivy respectively). "Finally on July 27 in a
shallow Quercus myrtifolia pond we heard so many oak toads we looked for
eggs. We find single bars of 26 or 8 eggs rarely attached to sticks at the sur-
face, usually attached to grass blades .5 to i or 2 inches below the surface of
the water. The water is 1-3 inches deep. Other bars are attached to pine
needles. Most often grass is the point of attachment. Once in awhile two
bars extend out from a common focus. Normally they are close together.
Found a female dead. Did she die from ovulation or mating?"

Detailed measurements of preserved material by Mrs. Wright give the
following results: 52 vitelli yielded 3 at .8 mm., 2 at .9 mm., and 47 at i.o
mm., the average and the mode i.o mm.; 51 tubes of jelly gave 23 at 1.2 mm.
in width, and 28 at 1.4 mm., the average 1.3 mm., the mode 1.4, but not one
actually at 1.3 mm.; 4, 5 or 6 eggs within a tube, each egg in sphere, spheres
stuck together, so partition between vitelli; of two 4-egg bars one is 4 mm.
long, one 4.8 mm. ; of two s-egg bars each is 5 mm. long; of one 6-egg bar it is
6 mm. long. In two accessions of eggs she notes "Queer little yellow drops
like drops of oil within egg tube and on outside". "Few drops Hke drops of
oil".

Dangers. In 1920 (p. 30) we held that "Many of these eggs or the sub-
sequent tadpoles are dried up by the rapid evaporation of the very transient

128 Frogs of the Okefinokee Swamp

breeding pools". No species suffers more badly than the oak toad from its
choice of breeding places yet it is an abundant form. Two to six or eight
inches are its usual depths for egg deposition. Often we made such remarks
as these in regard to eggs "The loss must be frightful". "How does this species
keep itself at abundance"? "The eggs in these furrows cannot escape dry-
ing". "The development must be rapid or the eggs or tadpoles will get
caught".

HATCHING PERIOD

Our first record of ovulation in 192 1 came June 4 at 10:00 p.m. Some of
these in petri dishes hatched June 8 before 6:00 a.m. when we arose. This
means from 72-80 hours from ovulation to hatching or 3-3 1/3 days.

MATURE TADPOLE

Color description from life (July 7, 1921). General coloration of the body
is grayish olive, olive lake, grape green or ecru olive produced by close set dots
of lighter color over a blackish background. Very few small black spots on the
back. In one two legged specimen general color quaker drab or of the vinace-
ous group. There is a black spot over the nostril and one over each eye. The
belly is pale purplish vinaceous; the throat and mentum with no bright color.
Gill region with clusters of deep olive buff or the general colors above.

Tail. The upper crest much more heavily marked than in Bufo terrestris.
Lower crest is also more or less marked with black. Along the dorsum of the
muscular part of the tail are 6 or 7 black saddles. Along the ventral edge of
muscular part are small black clusters but not so prominent as dorsal
saddles. In a two-legged specimen alternation of dark and light on tail
musculature for 8 or 9 spots. Along the back of body proper the charac-
teristic paired spots (4 or 5 of them) appear.

Iris black and general colors above.

General remarks. In 1923 (a, p. 406) Wright and Wright has ''Bufo
quercicus — gray tadpoles, six or seven black saddles on musculature, heavily
marked upper crest, venter one mass of color". There are plenty of oak toads
in the Okefinokee region. We had difficulty getting mated pairs at first.
Then when the developmental series of check pairs in the camp were fairly
well advanced, a severe rain came, overflowed the pans and some Bufo
quercicus tadpoles and Bufo terrestris tadpoles became mixed and our checks
for Bufo quercicus were never brought to mature tadpoles in the camp there-
after. There is a possibility of uncertainty in some of our Bufo quercicus
tadpoles, hence the unequal treatment given the tadpoles of this species.

LARVAL PERIOD

Deckert (1914 b, No. 9, p. 2) holds "the metamorphosis is very rapid".
In 1 92 1 we have the journal note July 7 that ''Bufo quercicus eggs laid June 4
are approaching transformation. One with four legs today." This implies 33
days. In 1922 we took mature tadpoles on June 27 implying egg laying before
our entrance to the swamp June 13-14. In 1922 we recorded transformation

Bujo quercicus 129

July 13 and 18. The earliest record of eggs we have in 192 1 or 1922 is June 4.
The intervals from June 4 to July 13 or 18 are 39 and 44 days. A period of
33-44 days does not seem unreasonable for such a summer breeder and shallow
water inhabitant though our evidence is slender (see tadpoles General Re-
marks) . Since the above was written we find that we captured a pair July 3 .
They laid July 4. The tadpoles were raised in camp and some went through
to transformation by August 12 or 39 days after egg laying.

TRANSFORMATION

Period. In 192 1 we recorded approaching transformation July 7. In
1922, on "July 12 found Bufo quercicus transformed in the edge of a pond in
wet pine barrens." On "July 13 around Coat Bet (Hyla gratiosa) pond on its
south side found plenty of young toads (Bufo quercicus) transformed. They
were near the edge of the pond but in the moist pine barrens especially in
clear areas". On July 18 (1922) found a few transformed around the shallow
edges of Murray Bay (a sphagnum swamp overgrown with heaths and even
trees). On August 16, 1922, near Callahan, Fla., we found them transformed.
At Trader's Hill Pond (1/2 mile west) on July 3, 1922 we found them mated.
Eggs from these were carried to camp. The transformed toads from this
series transformed August 1 2 .

In 1922 we then have transformation records from July 12-August 16.

Size. Deckert (1914 b, No. 9, pp. 2, 3) is the first to record anything on this
phase. The metamorphosis he holds to be very rapid. Hence one would
expect little pigmies of transformed frogs. "The young toads, when fully
developed, are 1/4 inch or less in length. The adults measure: male, i inch;
female, i 1/8 or i 1/4 inch".

On July 18, 1922 we secured five little toads at Murray Bay, Folkston, Ga.,
measuring 7, 7.5, 7.5, 8, 8 mm. respectively. They had "no stripes down back,
were prickly all over". On August 16 near Callahan, Florida, we took one
8 mm. long. On July 30-3 1, 1922, we secured one on Chesser Island, 10.5 mm.
long. It is just past transformation. The note on it reads: "Makes me think
of B. dehilis at this size". On August 12, 1922 we succeeded in transforming
this species at 7 mm. in length. Our largest series came July 12, 1922. Some
of them measured as follows: 6, 6, 5, 7 (median stripe), 7 ,7, 7, 7.5, 8, 8, 8, 8, 8,
8.5, 9, 9 (stripe down back), 9.5, 10 mm. respectively. Some of the larger
were slightly past transformation. The very tuberculate under surface of
forefoot suffices to distinguish it from Bufo terrestris.

GROWTH

This species transforms at 7-8 mm. On July 30, 1922, on Chesser Island,
Ga., we secured one 10.5 mm. and the next day in the same place one 13.5 mm.
In 191 7 at LeRoy, Ala., we took 9 specimens 11, 12, 13, 14, 14.5, 15, 16, 16,
16 mm. respectively and one 21.5 mm., possibly two age groups.

In the swamp we have isolated records such as: July 9, 1922 one 16.5 mm.;
May 6, 1921, one 15.5 mm.; August 13, 1922, 14, 15, 16, 17 mm. examples.

130 Frogs of the Okefinokee Swa7np

When we entered the swamp in 191 2, May 28, we picked up at Mixon's
Ferry, Suwannee River, the following: i at 15 mm.; 4 at 16 mm.; i at 16.5
mm.; i at 17 mm.; 3 at 17.5 mm.; 2 at 18 mm.; 2 at 18.5 mm.; 2 at 19.5 mm.;
I at 20 mm., probably one age group. Shortly after we arrived in 192 1 we
secured May i, 192 1, two 16, 16.5 mm. and one 25.5 mm. 9 . Certainly two
groups. It would seem as if these May and early June (to June 12 or 15)
records from 11-20 mm. are toads which wintered over and are i year beyond
transformation. At times it seems as if this group might be broken from the 2
year olds at 18 mm., but some of the 19-20 or 21 mm. are doubtful. Almost
all our material is determinable as to sex from 19 mm. onward for males and
from 20.5 or 21 or 22 mm. for females. Possibly the one year olds — if males to
be are 11-18 mm., and if females to be 11-20 or 21 mm.

We have collections on which we take a 17 mm. unsexed specimen and
four from 24-28 mm. sex determinable or a 18 mm. unsexed— apparently two
clear cut groups. But on June 10, 192 1, we took specimens, an 18 mm.
unsexed and 19.5, 20 and 20.5 mm. males or June i, 192 1, two unsexed 16.5
and 18 mm. and three sexed individuals 19 mm. cf , 21.5 mm. 9 and 22 mm.
9 . Almost certainly as determined in previous paragraph the 16-18 mm.
immature forms and mature group 19-22 mm. (sex revealed externally) are
two groups.

Most of our big series are as follows :

June 16, 1922 9 from 20.5 mm. 9-26.5 mm. 9

14 from 24.5 mm. cf-27.5 mm. cf

July 3, 192 1 7 from 23. mm. cf-26.5 mm. 9

1922 9 from 22. mm. 0^^26.5 mm. cf

Because of records such as these one might think 28-32 mm. individuals 3
year olds but the evidence is slender. For example, on May 25, 192 1, we take
together a 19 mm.cf and two females 28 mm. and 29.5 mm. or June 4, 192 1, a
male 26 mm. and a female 30.5 mm., or July 12, 1922, two 26 mm. d^s, 26.5
mm.. 9 and one 30 mm. 9 , or June 30, 1922, two 19 mm. cf s, two 25 mm.
9 s and 26 mm. cT and one 28 mm. cf , or July i, 1922, a male 24 mm. and one
male 29 mm.

In conclusion we hold transformation to come at 7-8 mm., a mean at 7.5
mm.; one year old stage 10.5 or 11 mm.-i8 or 19 or 20 mm., or mean at 16.5
mm.; two year olds, 19 cf or 21 9-27.5 or 28 mm. (possibly to 30 or 32 mm.),
mean at 25 mm. There may be a three year old class from 27.5 or 28 mm.-
32 mm., with mean 29 mm., but this is not quite so certain.

FOOD

For this topic consult Dr. R. Kellogg's forthcoming economic studies of
Bufos. He examined our series and found some very interesting things in its
diet. Professors C. R. Crosby and S. C. Bishop in examining some of Dr.
Kellogg's oak toad material thought so highly of the oak toad as an insect
collector that they entitled one paper thus, "A New Genus and two new
Species of Spiders Collected by Bufo quercicus (Holbrook)."

Bufo quercicus 131

ENEMIES

In 191 2 we wrote that the common garter snake ("highland moccasin")
prefers toads, and the oak toad with the Southern toad receives more of its
attention than any other prey.

In 1922 we felt that the spreading adder (Heterodon) is equally appre-
ciative of oak toads as well as southern toads. The gopher frog has also been
given as inordinately fond of oak toads.

AUTUMNAL DISAPPEARANCE

Our first record for the swamp is April 25 about the earliest we entered the
swamp and our latest date August 26, 1922. After our 1912 trip we received
specimens of this species collected from July 1 5 to November i , but there is no
certainty they were taken at the latter date though we suspect it near the
period of the oak toad's retirement. We find in literature no record beyond
our August 26 though it must stay out for two or three months beyond this
date.

AFFINITIES

Holbrook who describes it spends most of this phase of his description to
proving it is not the young of a large species. "That this little animal is not
the young of any other species I am certain, for

"i. It cannot be the young of the Bufo lentiginosus, for the superciliary
ridges are not elevated in proportion, the upper jaw is not emarginate,
and with the young of that species I am well acquainted.
"2. It cannot be the young of the Bufo americanus, as that animal is not
found near Charleston, and, besides, it wants the characteristic spade-
like process to the foot.
"3. It cannot be the young of the Bufo erithronotus, for its whole form is
different. It is not half the size, nor are its toes half as extensively
webbed; it is, in fact, a distinct and adult animal, for I have seen the
male and female together, and have seen the female deposit her spawn,
even when confined in a glass vessel".
It is a natural tendency to wonder if it be not a young of some other Bufo
when one first encounters it but it is very distinct as comparisons of an adult
B. quercicus and a young of Bufo lentiginosus reveal in the plate given (Plate
fig. 00).

LeConte did not make any mistake about it but Cope made it a new form,
Chilophryne dialopha. Boulenger (1882, pp. 319, 309) retained Cope's Bufo
dialophus of Sandwich Islands (p. 319) and places Bufo quercicus Holbrook in
the synonymy of Bufo lentiginosus americanus. Cope led him astray and he,
Cope, (1889, p. 292) alludes to it thus "The redescription of the species by
myself was due to the omission of its characteristic peculiarities from extant
writings. The erroneous locality (Sandwich Islands) is one of the several
such errors, based on the incorrect labehng of the collections of J. H. Town-
send, to which the specimen belonged."

132 Frogs of the Okefinokee Swatnp

In 1889 Cope (p. 292) writes of it as follows: "In some specimens the
transverse posterior part of the fronto-parietal crest is broken up. It then
resembles the young of the Bufo lentiginosus, with which it has been supposed
to be identical by various authors. It, however, differs from this species in
the differently shaped parotoid glands, the thickened posterior parts of the
mandibles, and from all the subspecies, except the B. I. woodhousei, in the
shorter head. There is no doubt but that Dr. Holbrook was correct in re-
garding this as a distinct species of very small size."

On the basis of vocal sac of male alone Miss Dickerson (1906, pp. 89-91)
places Bufo quercicus with Bufo compactilis and Bufo cognatus in relation-
ships. Other characters may bear her out in this determination.

In appearance one might place Bufo quercicus with Bufo cognatus and
might call B. quercicus sl small edition of it. Of course, there may be some
relationship with Bufo debilis the other small toad of the U. S. A., but it is
more remote from it than from B. cognatus. In the males the vocal vesicle
is of the Bufo cognatus groups but so is also some others such as Bufo com-
pactilis. Its note is quite different from the above species. We ourselves
know first hand the eggs of Bufo compactilis, B. punctatus and B. quercicus.
We have no positive eggs of B. debilis or Bufo cognatus. Bufo punctatus may
lay single eggs black and white, or eggs in loose flat masses on bottom or
quite a film on the bottom. Bufo quercicus lays brown and cream colored eggs,
4 to 6 or 8 in a bar with little or no jelly. Bufo compactilis lays brown and
cream colored eggs in a file, one envelope wide and with little jelly. In known
eggs of the United States Bufos, Bufo quercicus approaches closest to Bufo
compactilis. Possibly when B. cognatus eggs are found they will appear to be
related in appearance to Bufo quercicus and Bufo compactilis. This is, how-
ever, conjecture. The tadpoles of Bufo quercicus in coloration are lighter than
many Bufo tadpoles and may approach the bicolored tadpoles of Bufo com-
pactilis (as yet we do not positively know Bufo debilis and Bufo cognatus
tadpoles) .

Quite reasonably it seems to be in the Bufo cognatus-compactilis-punc-
tatus (possibly debilis) assemblage.

BIBLIOGRAPHY

1907 Brimley, C. S. A Key to the Species of Frogs and Toads Liable to Occur in North
CaroUna. Journ. EHsha Mitchell Sci. See. Dec. i, 1907, Vol. 23, pp. 157, 159.

1908 . Notes on Life Zones in North Carolina. Journ. Elisha Mitchell Sci. Soc,

May, 1908, Vol. 24, pp. 17-19.

1909 . Some Notes on the Zoology of Lake Ellis, Craven County, North Carolina,

with Special Reference to Herpetology. Proc. Biol. Soc. Wash., June 25, 1909,
Vol. XXII, p. 133.

19 10 . Records of Some Reptiles and Batrachians from the Southern United

States. Proc. Biol. Soc. Wash. March 31, 1910, Vol. XXIII, p. 11.

1926 . Revised Key and List of the Amphibians and Reptiles of North Carolina.

Journ. Elisha Mitchell Sci. Soc, Oct., 1926, Vol. 42, p. 80.
1862 Cope, E. D. Proceedings Acad. Sci. Phila., 1862, p. 341.

1886 . Proceedings Am. Phil. Soc, 1886, p. 516.

1889 . The Batrachia of North America. U. S. Nat. Mus. Bull. No. 34, pp.

291, 292.
1925 Crosby, C. R. and S. C. Bishop. A New Genus and Two New Species of Spiders

Collected by Bufo quercicus (Holbrook) The Florida Entomologist. October

1925, Vol. IX, No. 3, pp. 33-36.

Bufo terrestris 133

1914 Deckert, R. F. List of Salientia from near Jacksonville, Florida. Copeia Feb. 14,

1914, No. 3, p. 3.
1914 Deckert, R. F. Further Notes on the Salientia of Jacksonville, Florida. Copeia

Aug. 29, 1914, No. 9, pp. 2, 3.

1842 DeKay, James E. Zoology of New York, or the New York Fauna Part III, p. 68.

1906 Dickerson, M. C. The Frog Book, pp 89-91.

1842 Holbrook, John E. North American Herpetology, 2nd Edition, Vol. V, pp. 13-15.

1849 Holbrook, John E. Batrachia of Georgia Appendix (p. 15) in George White's

Statistics of the State of Georgia. Savannah 1849.
1855 LeConte, John. Descriptive Catalogue of the Ranina of the United States. Proc.

Acad. Nat. Sci. Phila. Dec. 1855, Vol 7, p. 430.

1922 Loding, H. P. A Preliminary Catalogue of Alabama Amphibians and Reptiles.
Geol. Survey of Ala. Museum Paper No. 5, Ala. Mus. Nat. Hist. Sept. 1922, pp.
16, 17.

1895 Loennberg, Einar. Notes on Reptiles and Batrachians Collected in Florida in 1892

and 1893. Proc. U. S. Nat. Mus. 1894, Vol. XVII, p. 338.
1924 Myers, George S. Amphibians and Reptiles from Wilmington, N. C. Copeia June

30^ 1924, No. 131. p. 59.
1924 Schmidt, Karl P. A List of Amphibians and Reptiles Collected near Charleston,

S. C. Copeia July 15, 1924, No. 132, p. 68.

1923 Stejneger, L. and T. Barbour. A Check List of North American Amphibians and
Reptiles, 2nd Edition, p. 26.

1923 Viosca, Percy, Jr. An Ecological Study of the Cold Blooded Vertebrates of South-
eastern Louisiana. Copeia Feb. i, 1923, No. 115, p. 37.

1920 Wright, A. H. Frogs: Their Natural History and Utilization. Appendix VI. Rept.
U. S. Commissioner of Fisheries 1919, Doc. 888, p. 30.

1923 . The Salientia of the Okefinokee Swamp, Georgia. Copeia Feb. i, 1923,

No. 115, P- 34-

1926 Wright, A. H. The Vertebrate Life of the Okefinokee Swamp in Relation to the
Atlantic Coastal Plain. Ecology Jan. 1926, Vol. VII, p. 82.

1923 Wright, A. H., and A. A. Wright. The Tadpoles of the Frogs of Okefinokee Swamp,
Georgia. The Anatomical Record, Jan. 20, 1923, Vol. 24, p. 406.

1924 Wright, A. H., and A. A. Wright. A Key to the Eggs of the Salientia East of the
Mississippi River. American Naturalist, July- August 1924, Vol. LVIII, pp. 375-381 •

Bufo terrestris Bonnaterre

(PI. I, Fig. 3; V, Fig. 8; VIII, Figs. 5, 6; XII, Fig. 1 1 ; XV, Fig. 1 1 ; XVII, XXII, Text Fig. I, 2)

COMMON NAMES

Southern Toad. Carolina Toad. Grey Toad. Land-frog (Bartram). Land-
toad (Catesby). Latreille's Toad. "Charming Toad." "Hop Toad."

RANGE

Check list. Type locality: "La Caroline."

Range. The Carolinas to Florida, west to the Mississippi. — Stejneger &
Barbour Check List, 1923, p. 26.

Ranges along the seacoast from South Carolina to Florida, and following
the Gulf through Alabama and Mississippi. Scattered individuals are met
with across the mountains to South Carolina. — Girard 1854, p. 86.

Supplementary records. In 1922 Deckert (Copeia, Nov. 20, 1922, p. 88)
found "Bufo terrestris'' singing at Royal Palm Hammock, and in wet places
northeast to Homestead." In June 1922 C. S. Brimley (1923, p. 4) in North
Carolina "found (them) common at Laurel Hill, Scotland Co., June i and 2,
1922." Earlier he (1922, p. 48) records "Southern Toad {Bufo terrestris).
One taken at Fayetteville, May 28, 1920, and two more on June 4 and 6,
1 92 1. The common toad of this region (Raleigh), however, appears to be

134 Frogs of the Okefinokee Swamp

Bufo fowleriy In 1922 in Duval Co., Fla., Thomas Hallinan at Fort George
Island found several. Nieden (1923, p. 125) though he publishes in 1923 really
comes about to the year 19 14 in data so has little or no additional range
evidence. He gives Southern United States, North America (east of Texas).
K. P. Schmidt (1924, p. 68) records it at Mt. Pleasant, S. C, and Bufo
fowleri at the same place. In South Carolina Pickens, 1927, has B. ameri-
canus, B. fowleri and Bufo terrestris in South Carolina. In 1923 Viosca
(1923, p. 37) gives Bufo terrestris as one of the Southeastern part of Louisiana.
At Wilmington, N. C, George S. Myers (1924, p. 59) took "Bufo terrestris
Bonnaterre. Several." In North Carolina, C. S. Brimley (1925, p. 80)
credits to to Jares, Onslow, New Hanover, Cumberland and Scotland Coun-
ties, In 1926 (pp. 82, 83) we considered it one of the "nine species of frogs
which might be considered Lower Austral." "I.O. Bufo terrestris S. C. — Ga.
La." Earlier in 1896 (p. loii) Cope placed "Bufo lentiginosus lentiginosus
Shaw" amongst the "Species which the Floridan subregion shares with the
Austroriparian . . ." He also (p. 1007) gives it as a characteristic Aus-
troriparian species. In 1889, Cope (p. 290) holds "The B. I. lentiginosus is
confined to the Austroriparian region east of Texas, and all statements to the
contrary are based on error. It does not ascend the Mississippi Valley, so
far as is known." The next year 1890 H. Garman reports it from Southern
Illinois and Kentucky. Did he have B. fowleri or is B. terrestris in Kentucky
and Illinois? Miss Dickerson (1906, p. 39) also places it in the Austroriparian
region. Brimley (1910, p. 11) has it from Bay St. Louis, Miss, to Bellair and
Fort Meade, Fla., or Mimsville and Riceborough, Ga. Viosca (1926, p. 308)
has it among "a few coastal plain species whose range from Virginia or North
Carolina southward and westward to the Mississippi Valley seems fairly well
established."

Shaw (1802, p. 173) credits it to the Carolinas and Virginia. Harlan
(1825, p. 345 and 1835, p. loa) does not consider terrestris alone when he has
it inhabiting "the southern and middle states."

Local Okefinokee records. In 19 12 fourteen adult or half-grown toads were
secured and about 100 others in various stages of growth beyond transforma-
tion. Several adults taken from snake stomachs were used in this (19 12)
study besides a series of transformed individuals and numerous tadpoles.

This species we found generally on all the islands and all around the out-
skirts of the swamp. No enumeration of localities is necessary. It occurs
throughout the pine barrens and hammocks in fact any land habitat. In
breeding we find it in shallow water normally. One Okefinokee boy called
them Charming Toad. "Charming toad because it charms you, turns your
eyes right green."

GENERAL APPEARANCE

Shaw (1802, p. 173) held "This very much resembles the common toad
(Europe) in its general appearance, but has a smaller head and sharper
snout. . . . Its colour is a dusky brown, paler beneath or it is all over
mottled with minute blackish or dark brown spots: the irides of the eyes
are red."

Bujo terrestris 135

In 1826 (pp. 344, 345) and in 1835 (p. 109) Harlan gives the following:
"Char. — Above dark-brown, verrucose, with irregularly disposed fuscus, or
blackish spots, edged with white: beneath dirty white, granulated: sides pale,
spotted; legs barred, large oblong warts behind the eyes; a large blackish spot
posterior to the tympanum; head above canahculate; two tubercles on the
heel of each foot; a longitudinal, vertebral, shallow, groove. Length of the
body about 3 inches."

Holbrook (1842, Vol. V, p. 7) gives its characters as: "Head large; snout
obtuse; superciHary ridges greatly elevated and terminating posteriorly in a
knob; upper jaw emaginate, lower furnished with a hook in front; parotoid
glands large, reniform, and reaching below the tympanum to near the shoul-
der; tympanum large; vocal vesicle internal; body above warty, dusky brown,
with a tinge of yellow; beneath granulated, dirty yellowish- white. Length
3 inches." Cope 1889 (p. 278) holds its definition to be "Frontoparietal
crests divergent, produced into a knob behind the short post-orbitals ; super-
tympanic well developed; head 3.5 to 4 times in length."

Miss Dickerson (1926, p. 43) chooses for distinction "head long (three
and half times in total length); cranial crests greatly elevated and swollen
behind; underparts unspotted." In 1907 Brimley (p. 157) chooses "bony
ridges ending in a knob behind" for his key character. In 1926 he uses the
same with the added note of "color usually dark and breast unspotted."

In 1914 Deckert (No. 9, p. 2) says "The toad itself, however, is dif-
ferently (from B. americanus) built, the head being wider and higher, and the
arms and legs shorter and more delicate. The eyes, also, are larger, and the
enormous bony knobs on the large heads of some old females give them a sort
of resemblance to species of the tropical cystignathoid toads Cemtophrys.
Unlike the latter, our toads are gentle creatures, living their lives of usefulness
in our farms and gardens."

Boulenger (1882, pp. 309, 310) gave his Bufo lentiginosus musicus these
characters: ""Supraorbital ridges swollen behind and produced beyond the
angles of the postorbitals; subarticular tubercles generally simple, metatarsal
tubercles moderate."

COLORATION OF SPIRIT SPECIMENS (191 2)
General color from buckthorn brown to almost black. Bright colored
forms have a prominent light vertebral band or line and a prominent oblique
Ught band from form of parotoid to the groin. In darker specimens the
lateral bands disappear before the vertebral line does. Many of the specimens
have both absent. The under-parts of one half of the adult series have the
breast and sides of the belly with fine spots like Holbrook's figure ('42, V,
fig. 11) while the other half have unspotted venters.

STRUCTURAL CHARACTERS (WRITTEN IN 19 1 2)

Cranial crest prominent; canthus rostalis low; preorbital ridge not ex-
tending below the level of the lower rim of the eye; supraorbital ridges
divergent and ending in elevated inward curved knobs which are not united
across the middle of the head by a flat bony rim except in one specimen. In

136 Frogs of the Okefinokee Swamp

this specimen (No. 6575) the supraorbital crests are lower, the furrow between
them shallow and the two ends merge across the middle line and thus form
an ovate bony-like plate between the eyes with the apex at the nostrils and
the base on a line of the anterior ends of the parotoids. At right angles to the
front of the supraorbital knob a ridge extends to a point just opposite the
end of the eyelid; here the ridge sends a branch downward 1/3 of the way
in front of the tympanum and sends another ridge backward over the tym-
panum to the parotoid; tympanic disk distinct, the cephalic border more
sharply defined than the caudal edge. Parotoid in some narrow and elongate,
in others shorter; in some reniform, in others oval; upper jaw slightly to de-
cidedly emarginate; head smooth; upper eyelid with small warts; back and
hind limbs with fine warts ; a spinose group above the arm insertion ; forearm
with coarse granulations; under parts granulate; 3rd finger slightly longer than
the first; 2nd finger longer than the 4th; tongue elliptical.

MEASUREMENTS

(1912— 1914)

The measurements of 14 adults or half -grown toads are as follows: In

length they are 40-67.75 mm., average 56.75 mm.; the head including the

tympanum is 13.5-25 mm., average 18.8 mm., mode 20 mm., contained in

length of body 2.9-3.4 times usually slightly over 3 times; or head to rear

of cranial crest 3-3.8 in length of body; width of head 15.5-28 mm., average

23.7 mm., mode 23 mm., usually slightly greater than head length, femur or

tibia; snout 5.75-8.75 mm., average 7 mm., mode 6 mm., equal to eye which is

5.5-8.25 mm., average 6.9, mode 6 mm.; tympanum 3.25-5.5 mm., average

4.3 mm., mode 4 mm., 1.2-2.0 in eye, usually at least 1.5 times; tympanum

slightly less than eyelid which is 3 . 2 5-6 . 2 5 mm . , average 5 mm . , mode 4 . 5 mm . ;

equals the interorbital distance or occasionally is slightly less, the interor-

bital distance being 4.25-6.25 mm., average 5.1 mm., mode 5 mm.; femur

13.5-29.5 mm., average 21 mm., mode 22 mm., equal to or slightly less than

rest of the foot, equal to or slightly more than the tibia which is 14-27 mm.,

average 20.5 mm.; rest of foot 15.5-29 mm., average 22 mm.; anterior limb

from axilla 26.75-44.5 mm., average 32.2 mm.; posterior limb from groin

48.5-78.5 mm., average 62.5 mm., or 1.7-2.3, usually under 2.0 times length

of anterior limb. ,^ . ^^ . . ,\

(Recent Material)

Head to angle of mouth 1.07 (20 mm.) 1.4 (44 mm.)-i.4 (56 mm.cf)

-1.37 (56 mm. 9)-i.33 (66 mm.cf)-i.35 (66 mm. 9)-i.4 (82 mm. 9 ) in

width of head; head to rear of tympanum ( ) — 1.3 — 1.16 — 1.26

— 1. 18 — 1.23 — 1.38 in width of head; head to angle of mouth 3.08 — 3.14

— 3.75 — 3.2 — 3.4 — 3.3 — 3.14 in length of body; head to rear of tympanum

( )-2.93 — 3.1 — 2.9 — 3.0 — 3.0 — 2.9 in length of body; snout i.o — .78

—.875^.875 — .7 — I.I— .96 in first finger; snout i.o — .78 — .875 — .875 — .75

— 1.0 — 1.0 in fourth finger; snout .66 — .50 — .437 — .437 — .50 — .66 — .50 in

first toe; eye 1.0 — 1.16 — 1.23 — 1.14 — 1.17 — 1.0 — i.i in snout; eye ( )

— .5 — .693 — .71 — .70 — .55 — .727 in tympanum; eye 1.0 — .91 — 1.08 — 1.0

— .82 — I.I — 1.04 in first finger; tympanum ( ) — 4.6 — 3.75 — 3.6 — 3.41

Bufo terrestris 137

—4.25 — 3.375 in intertympanic width; tympanum ( ) — 2.3 — 1.77 — 1.6

— 1.66 — 1.8 — 1.5 in snout; internasal width 1.33 — 1.3— 1. 12 — 1.25 — 1.3
— 1.3 — 1.08 in upper eyeHd width; interorbital width i.o — i.o — .9 — .83 — i.
— .92 — .72 in upper eyelid width; interorbital width .75 — .8 — .8 — .8 — .75
— .71 — .66 in internasal width; interorbital width ( ) — 3.0 — 3.4 — 3.0
— 3.4 — 3.0 — 3.0 in intertympanic width.

Forelimh. Forelimb 2.0 — 2.1 — 2.0 — 1.75 — 1.9 — 1.8 — 1.8 in length of
body; forelimb 2.2 — 2.15 — 2.1 — 1.7 — 2. — 1.8 — 1.85 in hindlimb; first finger
1.5 — 1.6 — 1.57 — 1.42 — 1.7 — 1.44 — 1.45 in third finger; second finger 1.5 —
1.65 — 1.83 — 1.42 — 1.7 — 1.44 — 1.45 in third finger; second finger 1.0 — 1.2
— 1. 16 — 1.0 — 1.0 — I.I — 1.45 in first finger; third finger .88 — .6 — .63 — .75
— .66 — .72 in second toe; fourth finger 1.5 — 1.5 — 1.55 — 1.42 — 1.6—1.55 — 1.3
in third finger; fourth finger .66 — .55 — .5 — .5 — .66 — .66 — .50 in first toe;
internasal width 2. — 1.6 — 1.75 — 1.75 — 1.5 — 2.0 — 1.9 in first finger; inter-
nasal width 2. — 1.4 — 1.5 — 1.75 — 1.55 — 1.8 — 1.82 in second finger; internasal
width 3. — 2.3 — 2.75 — 2.5 — 2.7 — 2.6 in third finger; internasal width 2. — 1.6
— 1.75 — 1.75 — 1.7 — 1.8 — 2.0 in fourth finger.

Hindlimb. Length i.i — 1.0 — 1.0 — 1.0 — 1.05 — 1.0 — 1.03 in hindlimb;
tibia 2.5—2.75—2.5—2.75—2.5—2.75—2.55 in length; tibia 2.75—2.8—
2.6 — 2.7 — 2.6 — 2.75 — 2.6 in hindlimb; tibia 1.25 — ^1.3 — 1.36 — 1.6 — 1.3 —
1.55 — 1.4 in forelimb; tibia .94 — .9 — .9 — .9 — .9 — 1.04 — i.oi in hind foot
(without tarsus); first toe 2. — 1.66 — 2.0 — 2.14 — 1.6 — 1.5 — 1.9 — in second toe;
first toe 3— 3— 3-5— 3-4— 2.8— 2.5— 3.3 in third toe; first toe 3.75—4.6—5.4
— 4.8 — 4.8 — 3.8 — 5.0 in fourth toe; first toe 2.5 — 2.8 — 3.1 — 2.9 — 2.8 — 2.8
— 3.0 in fifth toe; second toe 1.5 — 1.8 — 1.8 — 1.6 — 1.75 — 1.66— 1.75 in third
toe; second toe 1.87 — 2.9 — 2.7 — 2.26 — 3.0 — 2.5 — 2.6 in fourth toe; second toe
1.25 — 1.7 — 1.57 — 1.33 — 1.75 — 1.4 — 1.55 in fifth toe; third toe 1.25 — 1.6
1.5 — 1. 41 — 1.7 — 1.53 — 1.5 in fourth toe; third toe .83 — .94 — .9 — .83 — 1.0
— .83 — .9 — in fifth toe; fourth toe 1.3 — 1.5 — 1.6 — 1.7 — 1.55 — 1.6 — 1.55 in
fifth toe; fourth toe 1.6 — i.i — 1.15 — i.i — i.i — 1.05 — 1.06 in tibia; fourth
toe 1.33 — 1.45— 1.5 — 1.9 — 1.4 — 1.6 — 1.53 in forelimb; fifth toe 1.5 — 1.7 — 1.7
— 1.7 — 1.7 — 1.8— 1.7 in fourth toe; internasl with 1.3 — .9 — .9 — .9 — i.i
— 1.2 — 1.0 in first toe; internasal width 2.6 — 1.4 — 1.75 — 1.9 — 1.7 — 1.8 — 1.75
in second toe; internasal width 4. — 2.6 — 3.1 — 3. — 3. — 4.6 — 3.3 in third toe;
Internasal width 6. — 4.1 — 4.75 — 4.25 — 5.3 — 4.6 — 5.0 in fourth toe; internasal
width 3.3 — 2.4 — 2.75^ — 2.5—3.2 — 2.5 — 3.0 in fifth toe.

HABITAT

In 1892 and 1893 Einar Loennberg (1895, p. 338) finds "This is the most
common toad in south Florida. It is seen in great numbers about the houses
and other places in the evening. In the daytine, it hides under old logs,
boards, etc. at the shores of lakes and other moist places." Ten miles south of
Jacksonville, Florida, R. F. Deckert (Copeia, 1914, Feb. 14, No. 3, p. 3) re-
cords ''Bufo lentiginosus Shaw common in companies in temporary water or
scattered on cultivated ground.'" Again the same author (19 14, No. 9, p. 2)
writes "Bufo lentiginosus Shaw is the southern representative of our common

138 Frogs of the Okefinokee Swamp

(northern) toad, and seems to prefer the neighborhood of human habitations
for its abode. Near our house we found them under boards, logs, tubs, heaps
of rubbish, etc." In 192 1 he writes "It is found at any time of the year in
gardens and cultivated grounds."

(For habitat notes see General Habits, Ovulation Habitat, and Local Oke-
finokee records).

FIRST APPEARANCE

Most of the records for the first half of the year come in April. In the
U. S. National Museum collections we noted two collections in January,
three in February, four in March, eight in April, four in May, two in June
and eight in July.

In 192 1 we entered about April 22 and secured specimens at once. Two
days later they were breeding near our camp. This implies that they may have
been out much earlier.

GENERAL HABITS

Variations in color. Shaw (1802, p. 173) notes "They vary somewhat in
colour, being deeper or paler in different individuals."

Early naturalists in America began to find red toads and about them much
difference of opinion obtained.

Bartram (1791, p. 279) writes "The highland frogs, commonly called
toads, are of two species, the red and the black. The former, which is of a
reddish-brown or brick colour, is the largest, and may weigh upwards of one
pound when full grown; they have a disagreeable look, and when irritated,
they swell and raise themselves up on their four legs and croak, but are no
ways venomous or hurtful to man."

In 1827 J. L. Williams (A View of West Florida, 1827, p. 29) speaks of
"The red and black toads are common and useful reptiles; in destroying
insects they are extremely expert."

Harlan (1827, p. 345, 1835, p. 109) describes a "Var. A. The reddish-
brown, or brick-coloured toad of Bartram; very large, weighing near one
pound when full grown; legs and thighs marked with blotches and ringlets.
Inhabits the Southern States. The Red toad of Pennsylvania is but little
larger than the B. musicus."

Holbrook (1842, Vol. V, pp. 11, 12) figures and describes Bufo erythronotus
at considerable length. His last three topics might be repeated: "Habits.
This animal is found in the deep forests of oak in the neighbourhood of
Charleston, and is especially rare, as in nine years I have not seen more than a
dozen specimens, consequently nothing can be said of its habits.

"Geographical Distribution. As yet this animal has only been observed in
South Carolina.

"General Remarks. There is no doubt that this animal is an adult,
though I was at first disposed to consider it the young of the large toad of
similar colour mentioned by Bartram in his Travels in Florida, which as yet
I have never seen. Subsequent observation has, however, convinced me to
the contrary; for Bartram's animal weighed nearly a pound, while no specimen
of the Bufo erythronotus exceeded twenty lines in length; nor can it be the

Bnfo terrestris 139

young of Bufo lentiginosus, as the superciliary ridges are not elevated, and
the slight elevation that does exist is between the nostrils and orbits; or the
young of the Bufo americanus, which is sometimes red, for this latter animal
does not inhabit the low country of South Carolina, nor has it the spade-like
process at the foot, as in that animal; nor, in fact, is it the young of any
animal, as might be inferred from its size, for I have seen the male and female
together, and have seen their spawn."

In 1854 Charles Girard says little but it is direct. Under Bvfo lentiginosus
Holbrook he writes "Bufo erythronotus does not appear to us different from
B. lentiginosus, since color alone cannot be taken as an exclusive guide amongst
the bufonids."

In December, 1855, John LeConte (1856, p. 430) under Bufo musicus
Daudin writes "This species like all its congeners varies very much in color
and in the form and disposition of the spots. This variation depends en-
tirely on the will of the animal. It is generally, however, of a very dark
dusky, sometimes light grey and even reddish. I cannot conceive how it has
ever received the name of lentiginosus, which as I have shown before under
Telmatohius is as unlike this animal as any species can be. The Rana len-
tiginosa of Shaw, of which he gives in his general Zoology a tolerably good
figure, although copied from Catesby, is perfectly smooth and more resembling
a frog than a toad, and therefore well called by him the land frog; whereas
the B. musicus is very rough and makes no approach to its form and habits.
It walks rather than hops. Bartram's red toad of Florida was a mere variation
of this species, the difference being nothing more than an accidental or
voluntary change of color."

"Bufo erythronotus Holbrook, I.e. v. pi. 2. Not having been able for
years to obtain a living specimen of this animal, I shall say nothing more of it
than that it bears not the slightest resemblance to the Bufo lentiginosus
(B. musicus) as stated by Mr. Girard in Proceed. A. N. S., Vol. p. 86."

General habits. In 191 2 we write "This very abundant form is well known
and no account of its general habits is needed. We found it on the outskirts
of the swamp in the cut-over sections and it was common on the various
islands of the swamp, especially in the Lee's cornfields."

Shaw (1802, p. 173) speaks of its locomotion thus "Its motion also is not
that of crawling, but leaping." In general he says "These animals are said
to be most common in wet weather, but are very frequent on the higher
grounds, and appear in the hottest part of the day, as well as in the evening."
This is a paraphrase of Catesby. Harlan (1825, p. 345) holds it "Mostly
leaps, seldom crawls;" Holbrook (1842, Vol. V, p. 9) tells us "This animal
is timid and remarkably gentle in its habits, remaining concealed during the
day in some dark place, and only venturing out as the dusk of evening ap-
proaches." . . . "I have seen an individual kept for a long space of time,
which became perfectly tame: during the summer months he would retire to a
corner of the room, into a habitation he had prepared for himself, in a small
quantity of earth, placed there for his convenience. Towards evening he

I40 Frogs of the Okefinokee Swamp

would wander about the room in search of food, seizing greedily whatever
insect came his way. Some water having been squeezed from a sponge upon
his head one hot day in July, he returned the next day to the same spot, and
seemed well pleased with the repetition; nor did he fail during the extreme
heat of the summer to repair to it frequently, in search of his shower-bath."
In 1914 Deckert (No. 9, p. 2) holds "Their habits are in every respect similar
to those of Bufo americanus . . ."

VOICE

Holbrook (1842, Vol. V, p. 9) is the first who calls attention to its voice
''The males at this season are extremely noisy, though at other times they are
silent, or make only a sHght chirp when taken. Like many of the Hyla tribe,
they have a large sac under the throat, which is distended when the animal
croaks."

In 1908 H. A. Allard in discussing Bufo jowleri in Northern Georgia
(Gwinett & Jackson Counties) refers to the voice of Biijo lentiginosus thus:
"In this region throughout March I heard the occasional prolonged trills of
the so-considered common toad {Bufo lentiginosus). By the first of April
these notes had become quite silenced, and the distinct chorus of congrega-
tions of Fowler's toads had begun. I first noticed these toads singing on the
evening of March 26, although I think the first singers had appeared some-
what earher. It was interesting to note that the voice of Fowler's toads were
never heard with the appearance of cool, chilly nights, although the trilling
of the common toad continued. Throughout the early spring, this contrast
in the occurrence of the two notes, with respect to temperature changes was
very marked." (Science, 1908, n. s. 28, p. 655).

H. Garman (1890, p. 189) says that "The note of this variety is a singular
squawk which it is hardly possible to represent in words." Possibly the form
he had might have been B. foivleri. Deckert (1914, No. 9, p. 2) writes "the
call of the male cannot be distinguished from that of the Northern species."
In 192 1 he (Deckert) writes "this, the common Southern toad, was heard in
May, June, July, August and September, its trilhng call coming from rain-
ditches in hammocks, pineland and from roadsides."

In 192 1 we made these random notes on the first congress: April 24. The
note sounds more like our droning Bufo americanus, not Uke the scream of
Bufo fowleri. They are different .... Trill from 7 to 9 seconds in length,
shorter than Bufo americanus possibly.

When we entered the swamp April 22 the toads were well advanced in
their breeding period and less and less did they call in the day time unless a
heavy rain or downpour came. Like the meadow frogs they called more and
more in the evening and later and later in the evening as the season advanced.

The calling toads in cypress ponds and bays may be perched on a log,
on a cypress knee or stub or stump at the base of a cypress tree or gum tree,
on the moss, resting on aquatic plant stems, leaves, dead twigs usually in
shallow water or on the edge of a pond. In overflow pools it may be anywhere

Bujo terrestris 141

in shallow places. Rarely, if ever, does it float when croaking. In flooded
yam fields, cornfields, etc., it rests on the sides of the hills and may be a great
nuisance to other frogs (Hylas especially). Along the railroad ditches and
roadside pools it usually frequents the margins and edges when croaking.
If the water be not deep it is not particular as to stance, but it is truly an
alert, pert, "up on its toes" animal.

Some might think its note much like our Northern Bufo, some might call
it a trill, a drone, a musical note in quality or a bass roar. Many of them
nearby the observer are very deafening. Their choruses can be heard some
distance away.

The note is of 7-9 seconds duration, with intervals from 4-60 seconds.
Several counts Mr. Harper and I made were 14-23, 4i-5o> 4-1 1> i or 2 min.,
18-27, 45-55, 8-15, 28-38, ss-3, 16-24, 41-55, 10-7, 31-38, 52-60, or an
average of 31-55 seconds intervals. To Mr. Harper the "roar might be
represented thus: "waghrrrrrr rrrrrr." Just like other species they may
give weak notes; individuals may be freakish, hesitant, shrill or rarely open
their mouth to scream or half inflate their throats to give puzzhng notes.
Usually when croaking the throat is distended to its full capacity with com-
pressed body. The body is distended and the throat collapsed.

The voice record for 192 1 is as follows:

1921
April 23. Toads and southern meadow frogs began croaking in chorus in cypress

ponds 1-4 mile southeast of camp. Air is 58°, water 60°. Began about 8:00

or 9:00 p. m.
April 14. These began last night about 8:00. Air 66° at 9:15 a. m. Sun very

bright Moonshine Pond. Plenty of toads calling later than 7 :3o p. m.

" 25, Bufo in chorus a. m.

" 30. Tonight the air is cold. The thermometer at 7:00-8:00 p. m. is 58°.

Over at Long Pond and elsewhere we hear no Bufo terrestris. Possibly

because they are finished laying.
May 9. A lone Bufo terrestris calling. Last night went out at 8:30 p. m.

temperature 73°. Toads in bay at Billy's Lake were calling loudly. Could

hear no toads in other ponds.

" 15. Bufo calling at 9:00 p. m., temperature 65°. A cool wind and rain

from the east.
May 16. Many calling at night. Temperature 71°.

" 20. Heard several.

" 30. Heard a few Bufo terrestris. We need a rain.
June 4. Downpour. In Bufo pond a few heard. In Crosby pond heard no end

of them in deeper water. Temperature 70°. Hear plenty in negrao quarters.

In Bufo pond more calling than when I went out tonight.

" 5. Calling all about in rain puddles in cultivated fields at night.

" 17. 5w/o ^erresim calling in evening.

" 21. Late in night Bufo terrestris trifling.

" 22. Calling at 2:00 a. m. Temperature 88°.

" 23. Heard a few around Long Pond.

" 24. Heard one calling at night.

" 29. Commonly heard.
July I. Some toads going strongly.

" 3. No end of Bufo calling after the downpour. Temperature 72°.

142 Frogs of the Okefinokee Swamp

Jul}^ 4. Evening at 9:00 p. m. 75°, sultry. Toads are calling.

" 6. Abundantly calling about 10:00 p. m.

" 23. Last night I heard a few toads.

" 24. About two or three calling near camp at night.

July 26. Last night many toads calling.

" 28. Several calling about camp.

" 29. Bufo terrestris were commonly calling.

" 30. Toads calling last night.

Aug. 10. Several heard.

" 13. One heard.

" 14. Several heard.

" 16. Several calling at Camp Pinckney.

1922
June 19. Commonly calling at night in first dreen or slough.
July 2. Assembly Starling Branch 8:30 p. m., many calling.

" 3. Tonight from 8:00 p. m. to 1:00 a. m., July 4, heard toads. Some at

Trader's Hill Ponds. In many places they were heard.

" II. At second "dreen" several calling. A few heard at other places.

Temperature 70°.

" 25. Hard rain. Plenty of toads to be heard in the woods to the west edge

of the island.
Aug. I. Several in gutters of Waycross. Weak voice not deafening chorus.

" 2. One calling at night.

" 8. A few lone voices 8:00-11:00 p. m., after a rain. Temperature 79°.
Aug. II. Heard some 8:30-10:00 p. m., but rather few. In overflowed area of

Spanish Creek only chorus.

" 13. A few in road from St. Mary's to camp. In one or two instances a

fair chorus.

" 18. Several calling in evening.

" 19. One calling at night.

" 26. Several heard about dark.

Considering 12 choruses from April 23 to July 26, 192 1, we have for nearby
Weather Bureau Stations maxima 72°-94°, usual range 8i°-89°, average 84°,
mode 85°, for the day before; and minima from so°--j6° (one 45°), average
61°; for the day of the record, maxima from 8o°-92°, average 86°, mode 86°;
minima of 5o°-74°, average 66°. Heavy rains of .90-2.80 preceded most of
the choruses or at the time of the chorus. In the last of April maxima of
8i°-89° and minima of so°-62° obtained during croaking whether in chorus or
scattering calls. In the first half of May the maxima 72°-92°, minima 53°-67°
obtained. In the last of May maxima of 8o°-97° and minima of 63°-72°
obtained. In the first half of June maxima were 83°-9o°, minima 56°-7i°.
In the last of June maxima were from 8o°-98°, minima 67°-78°. In the first
of July maxima were 85°-92°, minima 67°-74°. In late July maxima were
85°-90°, minima 67°-72°. Thus we have minima of 5o°-62°, S3°-67°, 63°-72°,
56°-7i°, 67°-78°, 67°-74°, 67°-72°. The later the season of chorus quite
Ukely the greater the rain to start it.

The actual records of air temperatures taken by us in camp in 192 1 when
we heard Bitfo terrestris were 58°-88°. Most of the records range from
65°-75° and were preceded or accompanied by rain, or cloudy weather. When
we first entered the swamp on April 22 and 23 we had considerable rain.

Plate XXII

Southern toad {Bufo terrestris)

1. Bufo pond, Billy Id., Ga. June 6, 1921.

2. Female and male, Chesser Id., Ga. July 4, 1922. Ventral aspect, x 0.6.

3. Male croaking, Billy Id., Ga. June 4, 1921. Flashlight, x 0.3.

4. Male croaking, Billy Id., Ga. June 29, 192 1. Flashlight, x 0.3.

5. Male croaking, Billy Id., Ga. May 16, 1921. Flashlight, x 0.3.

6. Male (darkest) and two females, Billy Id., Ga. May 18, 192 1. x 0.3.

7. Bufo quercicus (adult) and Bufo terrestris (same size). Chesser Id., Ga. August

I, 1922. X I.e.

8. 9. Tadpoles, Camp Pinckney, Ga. July 24, 1922. x 1.0.

10. Transformed toad, Trail Ridge, Folkston, Ga. June 30, 1922. Dorsal aspect, x i.o.

11. Transformed toad. Trader's Hill, Ga. June 24, 1922. Lateral aspect, x 1.0.

12. Egg strings, stained, Chesser Id., Ga. July 4, 1922. x 1.3.

^ j^ t

• • • •♦ •

• ••- • T Z • •

#. ••••••

12

10

Biifo terrestris 143

The species bred from April 23-25. We heard them at 6o°-74°. On April 30
we had air 58° or less, in fact it went down to 50° or 5 1°. That night we heard
no toads from 5o°-58°.

In 1922 whenever we had a chorus or considerable calling from July 2 to
August 18 downpours, rain or cloudy weather obtained. Our lowest tem-
peratures during the day were from 7o°-8i°. The minima for nearby stations
were from 62°-74°. The best choruses are at least 68°-74°. Rain is the
important factor. Toads will call at 60° or lower but humidity dictates
largely in the matter.

MATING

Male {From life, June 9, 1921). Line from front line of eyes almost to vent
pale gull gray or mineral gray. Black spots along dorsum: one pair near
cephalic edge of eye ; one pair from upper eyelid connected across the meson ;
one in the mid dorsum between tympana; a pair either side of middle between
the rear ends of tympana; a pair of small spots; a pair of large spots where
hump comes; a pair of small spots; a pair just ahead of vent — all black spots
of dorsum thinly encircled with chalcedony yellow. More or less broken
pale gull gray or mineral gray line from tympanum to groin. Below this a
prominent black area. Tubercles of back black or deep brown tipped.
Lighter areas on rear of hind legs sulphur yellow. Either side of vent a few
orange tubercles. Under parts pale smoke gray. Pectoral region, under parts
of hind legs, and sides with black spots. Lower jaw rim like belly color.
Throat deep mouse gray or dark mouse gray with widely spaced white dots.
Gives throat discolored appearance. Light area of dorsum of hind foot same
color as the rear of thighs. Very little rusty on front of thighs and groin.
Top of first two fingers with excrescences and slight line of such on edge of
third finger. Color of excrescences chocolate or better hay's maroon or
maroon.

Female {From life, June 9, 1921). Lighter, larger. Practically no spots on
pectoral region. None on throat. Throat same color as belly. Practically
no rusty spots. Ground color of the dorsum more greenish olive. Some-
times at breeding season males and females may be alike in color, e.g., on
April 24, 1 92 1, several pairs were thus, several pairs reddish, one pair gray.
Most of pairs, however, were diverse.

Structural differences. Some of the differences noted in the field are :
I. On April 24, 192 1, we observed the following:

a. One male of a pair was about 3/5 of the female in size.

b. One pair, both male and female, were reddish. Two more pairs
red. One pair grey.

c. In some cases males and females alike in color but usually different.

d. One pair with one hind leg of female freshly eaten off, possibly by
a turtle.

e. The backs of the thumb and second finger of the male have ex-
crescences on the top and a slight excrescence on the inner edge of
the third finger.

1 44 Frogs of the Okefinokee Swamp

f . The females usually have the head crests larger, thicker and more
raised but this does not always obtain.
An examination of our and other collections reveal the following differ-
ence or notes:

a. The range of 84 males is 42-68 mm., one 82 mm., average 55 mm.,
mean 55 mm.

b. The range of 136 females is 44-88 mm., average 63 mm., mean
62 mm.

c. Usually the females have throat like rest of ymderparts and the
males have it discolored but some large females have throat clouded
(e.g., a 64 mm. 9 ) very clouded (e.g., a 76 mm. 9 ) or a male may
have throat quite light in color.

d. We have given 42-50 mm. as within the range of adult males.
As a matter of fact these have no excrescences on fingers or very
slight ones, but most of these have fully or partially discolored
throats. At about 54 mm. a male is sure to have excrescences and
discolored throat.

e. The heavy spotting in pectoral region may be in either sex or both.
The one median pectoral spot (like that of Bufo fowleri) occasion-
ally appears and may be in either sex,

f. Sometimes in the external recognition of sexes in the debatable
sizes of 42-49 mm. the higher or larger cranial crests in the females
help in the determination.

Duration. Usually the amplexation may last only an hour or so; if the
female be not ripe it may extend for days. Often the mated pairs in camp
might not lay the first night of capture. For example, "on July 2, 1922, at
Starling Branch Crossing 8:30 p. m. to midnight took one pair. Brought it
to camp. It did not lay on the night on July 2-3 and yet another taken at
same place July 3 from 8:00 p. m. to i :oo a. m., July 4, laid between i :oo-
7 :oo a. m., July 4, or 0-7 hours later."

On April 24, 192 1, two of the pairs were not broken by the next morning.
On May 2, we have a note about these that "the two toad pairs collected
April 23-24 are now broken," or 7 or 8 days later. On June 7 we have the
note that a pair "Have remained in embrace June 7 from night of June 4.
(Let them remain until June 9 when we forcibly broke them) "Or 5 days'
duration. Or on July 1 1 "at the second dreen (Chesser Id.) we found a mated
pair. . . Brought it into camp. . . . Temperature 70°. July 12. The
pair has not yet laid. July 16. The pair of July 11 fixed July 14."

C. S. Brimley in his notes on North Carolina Herpetology (Copeia, 1923,
p. 4) writes "Southern Toad {Bufo terrestris) found common at Laurel Hill,
Scotland Co., June i and 2, 1922. On June 2 a number were found breeding
on the edge of a roadside pool, and the following facts noted. The unmated
male toads at nine in the morning were screaming loudly on the edge of the
pool, but had not entered the water, while at noon they were in the water and
mated, but almost silent."

Bufo terrestris 145

Night or day.
April 23. Croaking in chorus in cypress ponds. Air 58°- water 60°. Began

about 8:00 or 9:00 p. m. Went over at 10:00 p. m. Caught two mated pairs of

Bufo. Took several males and one female. Brought these back. Mated two

males with two females, expect four pairs tomorrow morning.
April 24. The male and female of a pair if not yet laying are when taken out of

the water often given to urinating for protection. . . . Fifteen pairs in an

area 6 feet square. Sun very bright.
May 15. Rather cool. Toads are travelling everywhere.
May 16. Crosby Pond at night. In the pond are no end of Bufo terrestris. Several

mated pairs.
June 4. Everywhere in road home were toads. Hear plenty in lumber quarters.

It was 70° last night when I returned. In Bufo pond were more Bufo terrestris

than any night yet.
July 3. (In overflow area north of camp). Here in these temporary grassy

pools were no end of Bufo terrestris, pairs and single toads.

Amplexation. Normally the amplexation is axillary. The last two fingers
are not dug into the axil and often rest on the arm insertion of the female.

But, as with other toads, abnormal embraces are not unusual in this
species. On April 24, 192 1, we saw one lone male embracing the floating
rootstock of Pontederio. This male on the Pontederia root could be pulled
along or the root floated yet he would not let go.

Or, supra-axillary embraces may occur, e.g., June 7, 192 1, "The male
Bufo in camp of a mated pair is so small its embrace is supra-axillary. Had
dug into sides of female at least 1/3- to 1/2 inch above arm insertion almost
straight back from the tympanum and a little below middle of the parotoid."

Or subaxillary fashion may obtain. On July 3, 192 1, we have the note.
"In one mated pair of Bufo small male mating regular axillary fashion and
slightly back of axilla of small female. A large male holding a small female
behind fore hmbs and its hands on the breast of the female and almost
touching."

Occasionally a toad will attempt to embrace a meadow frog or leap at it.
When toads are breeding where Hyla gratiosa are, the impetuous toad males
will annoy the croaking Hylas considerably or any other Hyla or frog near
its size.

OVULATION

Habitat. In 192 1 we generally concluded that they preferred cypress
ponds, but the following notes will give some idea of some other habitats
found :

April 24. We captured some in recently dug post holes. After rain a night ago

these toads began to assemble in the ponds.
April 24. Moonshine pond. At its north end shallow and amongst the gums

plenty of toads calling.
May 9. Toads in cypress bay wooded swampy edge of island and also at

Billy's Lake where railroad spur goes across it.
May 16. Crosby pond. No end of toads. Some would float on surface of pond.

Many a toad would perch at base of a cypress tree or on a log or on a cypress

stub 3-8 inches above the water. Several mated pairs in water 4 inches to i

foot deep. Flashlighted only toads.

146 Frogs of the Okefinokee Swamp

May 31. No end of toads in temporary pools in negro quarters west.

June 4. Went to an open pond (no trees) south of our old camp. Here no end

of Bufo terrestris in deeper water.
June 6. Went to Oak Toad pond in cleared fields to photograph where oak toads

and Florida tree frogs were. Found plenty of Bufo terrestris eggs.
June 6. In temporary ponds in the street of the white quarters are toad's eggs.
July 3. In grassy overflow area toads common. Common in furrow pools of

newly planted sweet potato fields. In white quarters street pools no end of

Bufo terrestris.

Later we found them in roadside ditches, ditch pools beside the lumber
railroad, pools in tracks in wood roads, in dug pits, old field ponds, overflows
of branches, quiet waters of stream edges and countless other transient places.
On one occasion their eggs were so plentiful in impermanent pools that we
wrote June 4 ''What a frightful waste of frog life in transient pools."

Period. Holbrook (1842, Vol. V, p. 9) speaks of "the male (seeking)
the female in the month of May, when hundreds of them many be seen to-
gether in some stagnant pool, on which they deposit their spawn, and then
return again to the land."

During our stay 1912 a very few southern toads were heard and only one
record of eggs was made on Billy's Island, June 3, 19 12. These were lost
like those of Bvfo quercicus and were in similar situations. In the main, the
species doubtless is finished with egg-laying by the end of May, and it must
begin early for Mr. Harper reports "swarms of little toads appeared over the
paths, etc. about the middle of May, 191 2."

In 1 92 1 we have males croaking from April 23 to August 16, in 1922 from
June 19 to August 26. In 192 1 we have mating records from April 23 to
July 29 or 30; in 1922 from July 2 to August 13. In 192 1 we have ovulation
records from April 23 to July 3; in 1922 from June 22 to July 11. Through-
out my notes I have the ovulation period given variously as follows: April 15
or earlier to August i, August 13, August 17; April 24 to May 22; April 24
to August 13; April 24 or earlier to August i or August 15 or later; April i
to August 15 or later.

In 1923 (p. 34) we write "From April 15 or earlier to September i six
forms breed." Bufo terrestris is given as one of the six. In one set of notes
before we knew Pseudacris nigrita and Scaphiopus holbrookii from this region
we made it the third breeder after Acris gryllus and Rana sphenocephala.
It may be the fourth or fifth breeder if not the third in the sequence. In 1924
Wright & Wright (p. 380) give ovulation "Season, April 15 or earlier to
August 13."

In the light of the fact that most mating and ovulations were at night
from April 22 onward and because of transformation data as early as April
27 or mid May this species may begin as early as March i to April i or
earlier.

Temperature and humidity. On April 23, 192 1, we saw that toads would
breed at 58°-6o° or lower, but whenever ovulation occurred we generally had
records of 60° or higher, usually at least 64°-75°, the average 71°. In 1922
most of our records of air temperatures at ovulation were yi°-'jg° but the

Bufo terrestris 147

record runs farther into the summer. See Voice. As stated under voice a
drenching rain proves the important item in starting ovulation. We ob-
served that toads might breed at air temperatures of 58° or lower but 64°-; 5°
or 6o°^7i° seems a fair approximation of the necessary minimum. Rain,
however, is most important.

Egg-laying process. This species behaves in ovulation much as does Bufo
americanus. On April 24 we made a few observations. The unvarnished
journal notes are: When the female is not ovulating the male has the hind
legs free and floating, but when female ovulates several inches of egg string
the male brings his knees into the groin of the female and the heels almost
touch the upper surface of the hind feet against the under side of the female's
femur and near her cloacal opening. Female with hind legs stretched back,
sometimes heels touching, sometimes not. Eggs rest in cup made by heels
and feet of male. Male and female may remain in emission attitude 4 or 5
minutes or less. After the female crawls i foot or more, a minute or more
may elapse before another emission. When a female has begun ovulation it
it easy to pull the egg string out of the female. Water temperature where
laying 7 1°, air in sun 94°. Surface of water at 1 1 130 a. m. 81°.

On June 7, 192 1, we have the note that "Twice have Bufo terrestris
females laid a complement of eggs without the attendant male."

EGGS

Attachment. In many transient pools the egg strings are strewn on the
bottom or in some ponds in shallow water. More often the eggs are in strings
attached or woven about grass, vegetation, twigs or other support in the
water. In one pond we found them in a clear area and edge of a pond next
the Pontederia zone amongst scattering Sagittaria and another water plant.
These made a mat about which the eggs were woven. Sometimes they are
fastened near the surface in deep pools or along lake edges.

Egg mass, egg description. In 19 14 we wrote "Their characters if different
in minute form from those of B. I. americanus can not now be stated. In
general appearance, however, they were identical." Even in 192 1 we made
no detailed field notes on egg description but in 1922 on July 4 we have these
rough characters outhned. "Pair caught at Starling Branch crossing laid
between 1-7 a. m. today. Eggs seem to have only one jelly envelope. To-
night about 6:00 found a mass of eggs in Anna's pond. In road to Folkston
several masses found. The inner envelope of file revealed a day or more after
laying, not apparent when fresh. This inner envelope averages 2.0 mm. It
is near the outer envelope. No partitions between eggs. There is now a clear
elliptical envelope or membrane around each egg. These do not touch the
inner tube though. Possibly they are vitelline membranes. The outer
envelope is 2.8 mm."

On July II we caught a mated pair. On July 14 we fixed it and on July
16 we counted the eggs in the left ovary of the female. There were 1,444
eggs. This female was rather a small one, not over medium size at the most.
A total complement of 2,888 doubtless is small for this species.

148 Frogs of the Okefinokee Swamp

Detailed laboratory studies by Mrs. Wright are as follows. "July 4, 1922,
Starling Branch Bufo terrestris eggs. No inner envelope — no partitions — just
one homogeneous tube. 8-1 1 eggs in 30 mm. Black with small grayish-white
vegetative pole. Tube 2.8 mm. in diameter."

"Cannot find inner envelope or has inner tube swelled to meet outer tube.
Finally found inner tube but not very well marked. Inner tube present but
close to outer tube."

In another note we find "Tube inclined to be scalloped due to indentation
between each two eggs. Axis of developing vitellus longitudinal with tube."

Or "vitelli 2.4-2.6 or 2.8 mm. apart. In others 1.8-2.0 mm. apart."

Other notes are: "vitelli i.o, 1.2 or 1.4 mm., average 1.2, mean 1.2 mm.;
inner tube 2.2-3.4 mm.; outer tube 2.6-4.6 mm. 6 eggs in 20 mm.; 7 eggs
in 20 mm.; 7 eggs in 30 mm.; 8 eggs in 30 mm. and one 12 eggs in 30 mm."

Our final characterization in 1924 (p. 380) became the following: "Eggs in
files. Files long (several feet in length or often a meter or more long) ; vitellus
1.0 to 1.4 mm.; tube diameter 2.6 to 4.6 mm. Inner tube present. 7 or 8
eggs in 30 mm. (i 3/16 inches); distinct space between eggs; no partition
apparent; tube inclined to be slightly emarginate between the eggs; inner
tube 2.2 to 3.4 mm.; outer tube 2.6 to 4.6 mm. Egg complement, 2,888."

Dangers. If the eggs are laid in shallow water i to 4 or 5 inches from the
surface the danger of drying up in many ponds is grave to say nothing of the
transitory localities the species picks in periods of downpours.

HATCHING PERIOD

On April 24, 192 1, we found egg laying with water temperature 71°,
surface water 81° at 11:30 a. m. On April 28 we make the note "Bufo
lentiginosus eggs laid night of April 23-24 and morning of April 24 hatched
a day or so ago. Collected April 28, 4:00 p. m. Temperature 82°." This
implies 2 to 3 days for hatching. On June 22 found some eggs at Starling
Branch crossing presumably laid June 19. These were about ready to hatch.
They were in the open. But at Camp Pinckney in the shade on deeper water,
turpentine water pits they were not hatched though we make the note they
must have been laid June 19. These records would imply 3 or more days for
hatching. Our data is scanty. Hatching apparently comes in 2-4 days.

TADPOLE

Color description from life {July 7, 1921). General color is black with
scattered dots which are pale purplish vinaceous. Venter black with few
scattered light purplish vinaceous spots but not clustered to make one con-
tinuous mass of color as on the belly of B. quercicus.

Tail. Upper crest spotted but not so strongly as in Bufo quercicus. Lower
crest almost clear of spots. Lower edge of muscular part white or light chalce-
dony yellow.

General appearancQ. Body small (26 mm.) ovoid broader nearer vent than
eyes. Dorsal aspect shows body decidedly tapers ahead of the eyes. Dorsal

Bufo terrestris 149

crest low, extends but slightly on to the body to a vertical just ahead of the
buds of the hind legs. Dorsal and ventral crests about equal and about equal
to the depth of the musculature. Tail short, tip rounded. Spiracle sinistral,
directed mainly backward somewhat upwards, below the lateral axis. Eye
above lateral axis. Eyes dorsal, close together, slightly nearer lateral outline
than middorsal line. Anus median, opening in a level lower than the lower
edge of the ventral crest. Muciferous crypts indistinct.

Mouth parts: Teeth 2/3.

Upper labium fringed with a continuous row of labial teeth, the papillae
extending only each end of this fringe. The end of the second upper row does
not extend beyond the end of the upper fringe. The median space between .the
two parts of the second row is contained i. 4-2.1 times the lateral row Horny
beak about equal to or slightly less than the 3rd lower labial row of teeth
or contained 1.75 times in the upper fringe, or about 1.5 times the first or
second row of lower labial teeth, which are about equal. The third row of lower
labial teeth may be contained 1.2-1.4 times in the length of the second or first
lower lateral row. Rows of papillae at least two very prominent, sometimes
3-5 rows at corner of mouth. Lower loop of papillary border very prominent
with at least 2 distinct rows of papillae. This loop sometimes extends slightly
beneath the end of the third row of teeth. This loop decidedly lower in
level than end of the third labial lower row.

Measurements. Length of body (5.0-9.8 mm.) in tail (9.0-15.0 mm.)
1. 05-1. 66, average 1.3. Width (5.2-7.0 mm.) of body in its own length
1. 36-1. 74, average 1.51. Depth (3.6-5.0 mm.) of body i. 22-1. 6 in body
width, average 1.36 (2.4-5.0 mm.) of body 1.86-2.25 in body length, average
2.07. Depth of tail in length of tail 2.4-4.5, average 3.27. Muscular part (1.4
2.2 mm.) 1.5-3.0 in depth of tail, average 2.1 1. Spiracle 1.0-2.0 nearer base
of hind legs or vent region (3.6-6.0 mm.) than the tip of the snout (4.18-7.4
mm.), average 1.61. Spiracle i. 0-1.53 nearer eye (2.8-4.4 mm.) than base of
hind legs or vent, average 1.26. Eye i. 0-1.57 nearer to tip of snout (2.6-3.8
mm.) than to spiracle (2.8-4.4 mm.), average 1.19. Nostril 1.1-1.8 nearer
eye (i. 0-1.8 mm.) than snout (1.6-2.6 mm.), average 1.48. Mouth (2.0-2.8
mm.) usually 1.2-1.6 larger than internasal space (1.6-2.4 mm.), average 1.36.
Mouth contained i. 00-1.5 (average 1.17) in interorbital distance (2.0-3.4 mm.)
Internasal space contained in interorbital space 1.1-2.1, average 1.56.

The dimensions of the largest tadpoles are:

mm.

mm.

Total length

24

Spiracle to vent

4-5

Body length

9.8

Spiracle to eye

4.0

Body depth

50

Eye to snout

30

Body Width

7.0

Eye to nostril

1-4

Tail length

14.2

Nostril to snout

2. 2

Tail depth

4.8

Mouth

2.8

Musculature

of tail

1.6

Interorbital distance

3-4

Spiracle to snout

6.8

Internasal "

2.0

Measurements of 1912 specimens. The measurements of 20 specimens are
as follows: Length of body contained 1.2-2 times in the tail, average 1.37,

150 Frogs of the Okefinokee Swamp

mode 1.3. Width of the body in the length of the body 1.2-1.7, average
1.55. Nostrils .8-1.5 nearer the eye than the snout, average 1.2, mode 1.22,
one example 2.0. Eye lateral, usually much nearer to the snout than spiracle.
Distance between nostrils in interorbital distance .8-1.8 times, one example
2.0, average 1.25; in mouth 1.33-2.0, average 1.6. Spiracle 1.3 4-1. 7, usually
1. 4-1. 6, nearer the base of the hind legs than tip of snout, average 1.47. Anus
median. Depth of the tail in its own length 3.8-6.0, average 4.6. Depth of
the muscular part of the tail in the depth of the tail 1.0-2.5, average 1.5.
Interorbital distance is contained in the mouth i . 1-2 .0, average i .4. Greatest
length 20.25 mm. Greatest length of the body 8.5 mm. Greatest length of
tail 12.75 mm. Greatest depth of tail 3.0 mm.

General remarks. In coloration, mouth parts and measurements these
are not unlike those of Bnfo americanus.

LARVAL PERIOD

On June 22, 1922, we found eggs about ready to hatch in No. 2 turpentine
pit at Camp Pinckney. One July 25, at this same pit or 33 days later Bufo
tadpoles were transforming and transformed. We entered the swamp on
April 22, 1 92 1, on April 23 and 24 eggs were laid and our first transformation
records are April 27 and May 22. The latter record is probably not from the
April 23 and 24 eggs or if so it gives a month for larval period. The next trans-
formations come June i, June 4, June 16. These if from April 23 and 24
eggs would give 38, 39, 41, 42 and 54, and 55 days. Doubtless 38-55 days
falls within the period though less time down to 30 days may be probable
on special occasions.

TRANSFORMATION

Period. In 191 2 on June 12 in another more permanent pond a fine series
from tadpoles to transformations were taken. On June 24, still another
series of transformed frogs was taken.

In 192 1 on April 27 around Long Pond's edges we found tadpoles, trans-
formed toads and different sized young from 10 mm. to 24 mm. On May 7
in pools either side of the railroad found transforming young and different
stages of tadpoles. On May 15 many transformed young were travelling
on Billy's Island in the rain. On May 22 we secured more transformations.
On June i we note "toad tadpoles hopping around." On June 4 we make
three collections of transformed toads. On June 16 others were recorded and
transformations in July.

In 1922 we found southern toad toad tadpoles transformed at ponds i
mile south of Trader's Hill and also at Trader's Hill Landing on June 23.
Four days later we find a few in Anna's Pond. On June 30, Saddlebag Pond
has transformed toads. A month later on July 24 and 25 at Camp Pinckney
transformed and transforming toads were in evidence.

This gives us transformation from April 27 to July 25 but eggs laid in
August must transform in September or even October.

Size. In 191 2 the transformation sizes of 13 specimens range from 7.5-1 1.5
mm., average 9.75 mm., mode 9.5 mm. A series of June 24, 1912 was 6.5,

Bi(fo terrestris 151

7y 7-5) 7-5) 7-5) 7-5) 7-5> 7-5» 8, 8, 8, and 8.5 mm. In 192 1 our field notes
seemed to imply 8-10 mm. in size, but access to two series reveals the fol-
lowing: Several taken April 27 are 6.5, 7, 7, 7, 7, 7.5, 7.5, 7.5, 7.5, 8, 8, 8, 8, 8,
8, 8, 8, 8.5, 8.5, 8.5, 8.5, 9, 9, 9, 9, 9, 9-5, 9-5, 9-5, 1°, 1°, 10, 10.5 mm.; or five
of June 16 prove 8, 8, 8, 8.5, 9, 9.5 mm.

In 1922 on July 24 we secured a series of sizes as follows: 7, 7, 7.5, 7.5,
7-5, 7-5, 7-5, 7-5, 7-5, 7-5, 7-5, 7-5, 7-5, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8, 8,
8.5, 8.5, 8.5, 8.5, 8.5, 8.5, 8.5, 8.5, 8.5, 8.5, 9 transfg, 9, 9, 9.5. One specimen
appearing with this lot, is 12 mm. but it is far past transformation.

Apparently the range is about 6.5 — 10 — 10.5 mm., the mode 8 mm.,
average 8 mm.

GROWTH

If transformation comes at 6.5-1 1.5 mm., and males reach 84 mm. and
females 92 mm., how long a period elapses from emergence from tadpole
to the maximum sizes? Of if maturity comes at 42 mm. for male or 44 mm.
for female, but better about 50 mm. or more for each, what is the interval
between transfbrmation and maturity?

Ninety-six specimens (U. S. National Museum material, 7 lots) taken
in April by Dr. E. A. Mearns at Lake Kissimee seem to fall into the following
groups: 37-47 mm.; 48-59 mm.; 59-68 mm.; 69-78 mm. Some other ma-
terial in U. S. National Museum (158 specimens) falls into the following pre-
sumptive lots: 16-25 mm.; 25-35 mm.; 35-47 mm.; 48-59 mm.; 60-68 mm.;
69-78 mm.; 79-85 or 88 mm. On the basis of the 191 2 Okefinokee material,
the material seems to fall into groups 13-18 mm.; 27-29 mm.; 43- mm.;
48-59.5 mm.; 61.5-68 mm.; 70- mm. Our 1921 and 1922 material shapes
as 6.5-10 mm.; 13-19 mm.; 21-28 mm.; 34-37 mm.; 44 mm.; 48-56.5 mm.;
58-66 mm.; 69-79.5 mm.; 81-86 mm.; 92 mm.

Apparently this species transforms at 6.5-10 or 11 mm.; at one year may
be 13-25 mm.; at two years 25-35 mm.; three-year-olds 34-47 mm.; 48-59
or 60 mm. for four-year-olds; 59-66 mm. for five-year-olds; 69-78 or 79 mm. for
six-year-olds; 79-86 or 88 mm. for seven-year-olds; 90 mm. eight years or more.

The first two years of presumptive growth may be interpreted wrongly.
This species breeds over a long period and so it will be hard to arrive at
presumptive growths without actual records of marked specimens, e.g., our
13-25 mm. may not be correct, for on June 16, 1921, we took transformation
sizes from 8-9.5 mm. and three individuals 17, 21 and 28 mm. Should first
year's growth be 17-28 mm. instead of 13-25 mm.? That we dare not say.

Transformation comes at 6.5-10 or 11 mm. When do the parotoids,
cranial crests and tympana appear? We have examined our material of
sizes from 11-35 mm., one 12.5 mm. specimen taken May 22, 1912, ''ap-
parently past transformation" has none of these three structures. A series
of seven specimens taken May 31, 192 1, and ranging from 13-16 mm. have
not the tympana, crests or parotoid definitely outhned. In 1912 (May 15-
June 24) fourteen different accessions of material from 11-16 mm. in length
reveal none of these structures except two lots of 16 mm. One 16 mm. indi-
vidual had them not and two had no crests nor tympana but the parotoid

152 Frogs of the Okefinokee Swamp

were just outlined. All the accessions from 17 mm. onward had the parotoid
in different degrees of distinctness. From 11 to 22 mm. no tympana appear.
At 23 or 24 mm, the anterior half begins to appear as distinct but individuals
appear up to 24.5 mm. with no tympana. Some there are which do not have
it distinct until they reach 27.5-28 mm. To 21-23 mm. no cranial crests
appear but occasionally in this range crests are just outlined posteriorly.
From 23 to 24 mm. and onward the crests may appear as a groove or first
apparent posteriorly or as a fine row of tubercles. Occasionally a toad to
28 or 29 mm. may not have the crests outlined. At 34 or 35-37 mm. the
crests are beginning to thicken at their rear ends.

FOOD

Catesby makes the following notes "The Land Frog. The Back and
Upper part of this frog is gray, and thick spotted with dark brown spots; the
Belly dusky white, and faintly spotted: The Irides of the eyes red. They vary
somewhat in Colour, some being more gray, others inclining to brown. Their
Bodies are large, resembhng more a Toad than a Frog, yet they do not crawl
as Toads do, but leap; they are seen most in wet weather, yet are very fre-
quent in the higher lands, and appear in the hottest time of the Day: They
feed on Insects, particularly of one kind, which the following Accident seems
to confirm. As I was sitting in a sultry evening, with some company without
Doors, one of us let fall from a Pipe of Tobacco some light burning ashes,
which was immediately catched up and swallowed by a Frog of this kind.
This put us upon tempting him with a red hot Wood Coal, not less than the
End of one's Finger, which he also swallowed greedily; thus afterwards I
always found one or other of them easily deceived in this manner, as I imagine,
be taking it to be a Cicindela, or Fire-Fly, which on hot nights are very
numerous in Virginia and Carolina, where also these Frogs abound. — Catesby.
Vol. II, p. 69.

Shaw (1802, p. 173) writes "This animal is common in Carolina and Vir-
ginia, feeding, like most others of this genus, on insects, and is said to be
particularly attracted by any luminous insects, as fire-flies, glow-worms,
etc., and will even seize and swallow a small, live wood coal of the size of the
end of the finger; mistaking it for some luminous insect, and seeming to re-
ceive no immediate injury in consequence." Harlan (1825, p. 345) has it
"preying on living insects only." In 1842, Holbrook (Vol. V, p. 9) says "It
feeds on various insects which it seizes only while alive and in motion."

ENEMIES

In 19 1 2 these notes were made: The southern toad has a merciless set of
foes. The hog-nosed snake prefers them to all other diets and one snake may
have as many as two or three full grown toads in its stomach at one time. The
black snakes and garter snakes also are very partial to toads. In all instances
the head is swallowed first, and it is interesting to see how prominently the
head crests of B. terrestris stand out and how resistant they are to digestive
processes when the other forward parts of the body are well digested.

Bufo terrestris 153

In 192 1 we held the kilhfishes such as Gambusia, Lucania or Fundulus
doubtless did little damage to the eggs. Possibly some newly hatched young
might suffer. In some ponds turtles do considerable damage to tadpoles, or
to adults as well. In some of the breeding congresses we have chanced on
southern water snakes hunting for toads or frogs. At times we have caught
them in the act of seizing a toad or frog or with the subject partially down.
In 1922 on July 15 on our way out to Folkston we found a spreading adder.
It began to hiss and spread. By pressing a stick on it it turned over on its back.
By pressing farther it disarticulated its lower jaw and lolled out its tongue and
finally disgorged a freshly caught southern toad.

AUTUMNAL DISAPPEARANCE

The Cornell parties which entered the swamp in December and January
reported it in these months. In the U.S. National Museum collections there is
scant if any material taken in October and November and first of December.
In the same way our collection has little at these months. Doubtless from
November to January is its most inactive period.

AFFINITIES

In 191 2 we wrote ''Relationships. — The American toads of the old Bufo
lentiginosus series are yet in an unsatisfactory stage of analysis. At first Cope
employed the cranial crests as one of the cardinal characters of differentiation.
On this criterion Bufo terrestris and Bufo americanus can usually be separated,
the former with posterior ends of the frontoparietal crests very much elevated
and swollen and the latter with this region and whole crest low. Usually
this character suffices to separate the two though we can find within these
two a small degree of variability. The other two forms B. fowleri and B.
woodhousei have the crests parallel or divergent, raised or low, merged at
posterior ends or free and hence Miss Dickerson discards the character and
ascribes the confusion within this assemblage almost solely to the reliance on
this one character. If we rely on color we encounter an almost perfect B.
terrestris duplicate (No. 6575) of the B. fowleri in the vertebral arrangement
of the six pairs of dorsal spots as given by Miss Dickerson (p. 64). To be sure
the writers' impressions of the color of B. terrestris and B. fowleri are that the
former averages darker than the latter while the latter inclines towards the
greenish or yellowish grey. The under-parts of Bufo terrestris are not always
unspotted and Holbrook's figure shows the reverse condition. Our specimens
are about evenly divided on the point, and rarely B. fowleri from N. J.,
L. I., and Woods Hole, Mass., are somewhat spotted though not as markedly
so as in the northeastern Canadian specimens of B. americanus while some of
the B. americanus of Ithaca may be practically if not solely unspotted. The
head length is equally emphasized as a good distinction but since Cope's
time most authors have not defined what they mean by head length in toad
descriptions. Cope measured to the end of the frontoparietal crests and Miss
Dickerson in her figure of terms employed gives the usual measurement to the
caudal border of the tympanum. On the latter basis some B. fowleri would

154 Frogs of the Okefinokee Swamp

have heads 3.5 in the length while obviously her 4 to 4.5 times in the body
refers to Cope's method of measurement. Some of B. ayyiericanus according
to Cope's method would be 4 in the body, but if measured to the tympanum
would be 3.5 in the length. Or a Texan B. woodhousei with head 4.6 in the
length by cranial crests test would be 3.5 by the tympanic rule, quite far
moved from "head five times in total length." The double file noted in the
egg strings of the European Bufo vulgaris, has been urged as distinctive of B.
fowleri while B. americanus never has such an arrangement. Rarely, B.
americanus eggs are in the double file and certainly B. americanus can and
does breed from April 20-August. The author has seen the eggs of Bufo
fowleri in two instances only and in each case they were more or less a double file.

"At the present, most beginners and some experts, some of which are with-
holding judgment, are unable to determine whether B. fowleri is distinct
enough to be considered a separate species whether it exists as a distinct form
at all or whether it needs an even finer refinement of descriptive characteristics.
At present the authors withhold judgment and merely show some of the
difficulties. B. americanus and B. lentiginosus seem quite distinct. Of B.
fowleri we say it probably exists as a separate form but needs description with
an abundant comparative series from Ontario to Texas, from which regions
our series has been taken but not in sufficient numbers to warrant a decision."

The foregoing paragraphs were written in 1913 but today the author is not
yet ready to speak any ill-considered conclusions on the Bufo americanus-
fowleri-terrestris-woodhousii complex. I have not worked on the life history
of B. woodhousii, have seen eggs of Bufo fowleri but four times and need more
personal experience with these two species. In its egg characters Bufo
terrestris has two jelly tubes like B. americanus but like Bufo fowleri may have
no partitions between the eggs. We have tadpoles of Bufo americanus, Bufo
terrestris and Bufo fowleri but feel hesitant to venture opinions on their degree
of relationship. On superficial morphological characters, in some respects,
Bufo terrestris appears as much related to Bufo fowleri as Bufo americanus.

Holbrook (1842, Vol. V, p. 10) reviews its history to 1842. We give his
"General Remarks. Catesby first described and gave a figure of this animal
under the name of Land-frog; and although his figure is badly executed, both
as to drawing and colouring (the elevation of the superciliary ridges not being
marked, and the eyes represented as red) , it has been repeatedly copied by later
naturalists, as Foster, Shaw, etc. The name, however, Rana (Bufo) terrestris,
cannot be retained, as it is previously applied to another animal.

"Bosc, who from a long residence in Carolina, had a good opportunity of
examining this animal, refers it to the Rana musica of Linnaeus, in which he
is followed by Daudin, Merrem, and most naturalists. This cannot be correct,
for there are no toads, as far as has been hitherto ascertained, common to
North and South America; and Linnaeus, in the twelfth edition of the Systema
Naturae, gives Surinam, as the country of his Rana musica. Neither the
specific name, terrestris, nor musicus can then be applied to this animal, but
we must give it the one next in order under which it is found described —
Rana (bufo) lentiginosa of Shaw."

Bufo terrestris

155

In 1849 (Appendix p. 15) Holbrook gives both Bufo lentiginosus Shaw and
Bufo erythronotus Holbrook as of Georgia. In 1854 Girard (p. 56, p. 86) Usts
Bufo americanus LeConte, Bufo lentiginosus Holbrook with Bufo erythronotus
as a synonym, Bufo woodhousii Girard, Bufo cognatus Say, etc. He was
nearer the situation than LeConte in 1855 (1856, p. 430) who gives B. musicus
Daudin {B. lentiginosus Holbrook), Bufo americanus, B. erythronotus. His
comment that Girard is wrong in referring B. erythronotus to Bufo lentiginosus
may be on a par with his placing Bufo cognatus in Bufo muscicus. DeKay
(1842, p. 68) gives both Bufo lentiginosus and Bufo erythronotus each distinct,
also Bufo cognatus as distinct and all three as extralimital to New York State.
Only Bufo americanus does he credit to New York.

In 1858 Albert Gunther (p. 63) gives Bufo lentiginosus as consisting of two
varieties "Var. A. Bufo americanus .... Bony ridges moderate, but swollen
behind and "Var. B. Bufo musicus .... Bony ridge swollen behind.

In 1882 Boulenger (pp. 306-310) gives Bufo lentiginosus as having 5
subspecies, namely, Var. A. americanus (quercicus, woodhousii, etc.), Var. B.
musicus (lentiginosus), Var. C. frontosus, Var. D. cognatus, Var. E. fowler i.
His Var. B. musicus (lentiginosus) is with "Supraorbital ridges swollen behind,
and produced beyond the angle of the postorbitals; subarticular tubercles
generally simple; metatarsal tubercles moderate." Yarrow, 1882, (pp. 23,
164-167) gives the same five forms of Bufo lentiginosus. Cope (1886, pp.
515, 516) has Bufo cognatus separate from Bufo lentiginosus complex and
describes Bufo hemiophrys. His B. lentiginosus he has as "B. 1. fowleri ....
Canadian and Hudsonian districts of eastern region." Bufo 1. woodhouseii
. . . Central region. Bufo 1. americanus . . . eastern and Austroriparian
regions." "Bufo 1. lentiginosus . . . Austroriparian region."

In 1896 Rhoads (p. 396) in Tennessee took "Bufo lentiginosus (Shaw)
Southern Toad. Specimens from Southern Tennessee approach nearly to
those found in the Gulf States but the majority are intermediates ....
On Roan Mt. he found B. lentiginosus americanus at 5,000-6,300 feet.

BIBLIOGRAPHY

1908 Allard, H. C. Bufo fowleri (Putnam) in Northern Georgia. Science N. S. Vol. 28,

No. 723, Nov. 6, 1908, pp. 655, 656.
1 79 1 Bartram, Wm. Travels through North and South Carolina, Georgia, East and West

Florida. Phila, 1791, pp. 279, 280.
1882 Boulenger, G. A. Catalogue of the Batrachia Salientia S. Ecaudata . . . British

Museum 2nd edit. London, 1882, pp. 309, 310.
1907 Brimley, C. S. A Key to the Species of Frogs and Toads Liable to Occur in North

Carolina. Journal Elisha Mitchell Sci. Soc, Dec. 1907, Vol. XXIII, p. 157.
19 10 Brimley, C. S. Records of some Reptiles and Batrachians from the Southeastern

United States. Proc. Biol. Soc. Wash., Mar. 23, 1910, Vol. XXIII, p. 11.
1922 Brimley, C. S. Herpetological Notes from North Carolina. Copeia No. 107, June

20, 1922, p. 48.

1923 : North Carohna Herpetology. Copeia, Jan. 20, 1923, No. 114, p. 4.

1926 . Revised Key and List of the Amphibians and Reptiles of North Carolina.

Journal Elisha Mitchell Society, Vol. 42, Nos. i and 2, October, 1926, p. 80.
1886 Cope, E. D. Synonymic List of North American Species of Bufo and Rana,

Am. Phil. Soc. Proc, 1886, Vol. 23, pp. 515, 516.

1889 . The Batrachia of North America. U. S. N. Mus. Bull. 24, 1889, p. 290.

1896 Cope, E. D. The Geographical Distribution of Batrachia and Reptilia in North

America. The American Naturalist, Dec. 1896, Vol. 30, p. 1007.
1914 Deckert, R. F. List of Salientia from near Jacksonville, Florida. Copeia, Feb. 14,

No. 3, p. 3.

156 Frogs of the Okefinokee Swamp

19 14 Deckert, R. F. Further Notes on the Salientia of Jacksonville, Florida. Copeia,
August 29, 1914, No. 9, p. 2.

1921 Deckert, R. F. Amphibian Notes from Dade Co., Florida. Copeia, Mar. 15, 1921,
No. 92, p. 21.

1922 Deckert, R. F. Notes on Dade Co. Sahentia. Copeia, Nov. 20, 1922, No. 112,
p. 88.

1842 DeKay, J. E. Natural History of New York, Part III, Albany, 1842, p. 68.
1906 Dickerson, M. C. The Frog Book, pp. 39, 45.

1890 Garman, H. Notes on Illinois Reptiles and Amphibians .... Bull. 111. State Lab.

Nat. Hist., Vol. Ill, Peoria 1896 Art X, Sept. 1890, pp. 189, 190.
1854 Girard, Charles. A List of the North American Bufonids with Diagnoses of New

Species. Proc. Acad. Nat. Sciences Phila., Vol. VII, 1854, 1855. Phila. 1856, p. 86.
1827 Harlan, Richard. Genera of North American Reptilia and a Synopsis of the Species.

Dec. 12, 1826. Journal Acad. Nat. Sci. Phila., Vol. V, Part ll, 1827, pp. 344, 345.
1835 . Medical and Physical Researches, pp. 109, no.

1843 Holbrook, J. E. North American Herpetology, 2nd Edition, Vol. V, pp. 7-12.
1848 . Reptiles. In Statistics of Georgia by George White, Savannah, 1849,

Appendix p. 15.
1856 LeConte, John. Descriptive Catalogue of the Ranina of the United States. Proc.

Acad. Nat. Sci. Phila., Vol. VII, 1854, 1855, 1856, p. 430. (Dec. 1855).
1895 Loennberg, E. Notes on Reptiles and Batrachians collected in Florida in 1892 and

1893. Proc. U. S. Nat. Mus., Vol. XVII, 1894, Wash. 1895, p. 338.
1924 Myers, G. S. Amphibians and Reptiles from. Wilmington, N. C. Copeia, No. 131,

June 30, 1924, p. 59.

1923 Nieden, Fr. Das Tierreich. Anura I, Berhn and Leipzig, pp. 124, 125.

1927 Pickens, A. L. Amphibians of Upper South CaroUna. Copeia, Oct.-Dec. 1927.

No. 165, Dec. 23, 1927, p. 109.
1895 Rhoads, S. N. Contributions to the Zoology of Tennessee. No. i Reptiles and

Amphibians. Proc. Acad. Nat. Sci. Phila., Vol. 47 (1895), p. 396.

1924 Schmidt, K. P. A List of Amphibians and Reptiles. Collected near Charleston,
S. C. Copeia, July 15, 1924, No. 132, p. 68.

1802 Shaw, George. General Zoology Amphibia Vol. Ill, Part I, 1802, p. 173.

1924 Viosca, Percy, Jr. An Ecological Study of the Cold Blooded Vertebrates of South-
eastern Louisiana. Copeia, Feb. i, 1923. No. 115, p. 37.

1923 . Distributional Problems of the Cold Blooded Vertebrates of the Gulf

Coastal Plain. Ecology VII, pp. 307-314.

1827 Williams, J. L. A View of West Florida, p. 29.

1923 Wright, A. H. The Salientia of the Okefinokee Swamp, Georgia, Copeia, Feb. i ,
1923, No. 115, p. 34.

1926. Wright, A. H. The Vertebrate Life of the Okefinokee Swamp in Relation to the
Atlantic Coastal Plain. Ecology VII, No. i, January 1926, pp. 82, 83.

1924 Wright, A. H. and A. A. Wright. A Key to the Eggs of Salientia East of the Missis-
sippi River. The American Naturalist, Vol. LVIII, July- August, 1924, p. 380.

1883 Yarrow, H. C. Check List of North American Reptilia and Batrachia. U. S. Nat.
Mus. Bull. No. 24, 1883, pp. 23.

Acris gryllus (LeConte)

(PI. II, Fig. I ; V, Fig. 5; VI, Fig. i ; X, Fig. 1 1 ; XII, Fig. 6; XV, Fig. 7; XVII; XXIII; Text

Fig. I, 9)

COMMON NAMES

Cricket-Frog. Savannah Cricket. Savanna Cricket. The Cricket
Hylodes. Peeper. Rattler. Western Cricket Frog (crepitans). Savannah
Cricket Frog. Cricket Toad. Western Cricket (crepitans).

RANGE

Check list. Florida to New York in the east ; northward through the central
valley from Louisiana and Texas to the Canadian Northwest Territories.

Supplementary records. Cope 1892 (p. 333) had them from the Staked
Plains of Texas. Stone (1903, pp. 538-539) had them from Del Rio, Texas.
Strecker (1908, p. 58) has it from Borachio, El Paso County, Texas.

.4cm gryllus 157

In the western states Ellis (19 13, p. 58) reports it at Wray, Colorado and
in 19 1 7 (p. 39) he reports it from Roswell, New Mexico.

In the extreme northeast Babcock in 192 1 seriously questions the two
previous records of its presence in New England.

In the north we have records from Baltrusol, N. J. and Passaic River
Summit (A. C. Chandler 1909) to Hillsdale Michigan (T. L. Hankinson 1900)
to Ames, Iowa, (Mr. and Mrs. (K. Van Winkle) E. L. Palmer 1924) to Emporia
and Lawrence, Kansas (G. D. Hanna 1909). From Chapel Point, Maryland
(Harper 19 16) to Gloucester, North Carohna (Harper 19 13), and Raleigh
(Brimley, 1897) to Columbia, South Carolina (Corrington) to Caloosatcha
River (J. C. Bradley 191 1); we have it in our collections of the Atlantic Sea-
board. From Georgia through Bay St. Louis, Miss., (1897) to Doucette,
Texas (O. D. Ingalls (1909), it occurs in the Gulf States.

In 191 7 we collected it from Petersburg, Va., May 31 and June i to
Neuces River, Texas July 2, and the following intermediate localities: Din-
widdle, Va., June i; Soudan, Va., June 3; Flatwood, Ala., June 9; LeRoy,
Ala., June 12; Theodore, Ala., June 13; Biloxi, Miss., June 14; Dayton, Tex.,
June 22; New Braunfels, Tex., June 27. In countless places we recorded it
between Virginia and Texas.

In 1925 we went out to New Orleans, La., with P. L. Viosca, Jr., who
knows A. gryllus and A. crepitans better than any student of our Anurans.
In Texas from Feb. 8, to June 30, 1925, Mrs. Wright and I collected Acris
gryllus about San Antonio and at Helotes Creek, Texas. Southward we took
it at Beeville, Texas, March 25 to May i, 1925. Westward we found it 23
miles east of Del Rio, Texas, June 30, 1925.

In 1926 (p. 83) we termed "Acris gryllus'N.Y. — Fla. — Tex." as a Lower
Austral frog but it extends over into Upper Austral regions.

In Georgia, the State of which the Swamp is a part, from 1910-1911 Prof.
J. C. Bradley secured it as follows: Sept.-Oct., 1910, Spring Creek, Decatur
Co.; Mar. 5, 191 1, Marietta; March 8, Wrens; Apr. 2, Gainesville; Apr. 22,
Offerman, Satilla River; May 19, St. Simon Id., and June 7-23 Spring Creek,
Decatur Co. In Cuthbert, Ga., March 29, 192 1, F. Harper took this species.

Local Okefi.nokee records. In 19 14 we wrote: "Our material consists of a
series of tadpoles to transformation taken from May 28- June 15, 1912, and
of 44 specimens of frogs of which 38 were secured between the above dates, 3
taken by Mr. Harper from May 2-23, 1912, and 3 on Dec. 22, 1913. This
species was easily the commonest member of the Hylidae within the swamp.
Most of our specimens came from Billy's Island, but representatives of the
species were recorded or taken on Honey Island and its prairies, Floyd's
Island and its prairies, Minne Islands, and at Mixon's Ferry and Hammock."

In 192 1 we took it at Cypress Pond, Long Pond, Bullfrog Pond, Crosby
Pond and other points on Billy's Island, Chesser Island, Black Jack Island,
Honey Island and on prairies. Honey Island, Chase Prairie, and Black Jack
strand ; on Minne Lake River and a detour. We also recorded it in the Pocket ,
on Jones Island, Gallberry Island, Floyd's Island Prairie, Billy's Lake, and
almost all over the swamp.

158 Frogs of the Okefinokee Swamp

In 1922 we took it from Folkston to Jacksonville, Fla., from Folkston to
Chesser Island, Okefinokee Swamp, at Camp Cornelia, Starling Branch, St
Mary's River at many points and universally in the region surveyed. On the
trip from Ithaca, N. Y. to Okefinokee Swamp June 3-14, 1922, we first heard
Acris near Dyke Marsh, near New Alexandria, Va., and 12 miles north of
Petersburg, Va., on June 7. Near Cheraw, S. C, June 8, we heard them at
the same lake around which we heard Hyla andersoni and Rana virgatipes.
At Mabee, June 9, Acris was calling at noon. On June 10 at night camp near
Millen, Ga., they were calling and thus onward to Chesser Island, Okefinokee
Swamp.

GENERAL APPEARANCE

Its original description (LeConte, 1825, p. 282) is "Rana gryllus (Savanna
Cricket), above warty, colour various, with a triangular spot of darker on the
top of the head between the eyes, and a paler line extending from the apex of
this spot to the vent, hind part of the thighs yellowish or whitish, with one or
two lines of dusky or brown."

Holbrook (1842, p. 131) gave its outstanding characters as "Head elon-
gated, pointed, a triangular dusky spot between the orbits; body above
cinerous with a green or sometimes red, vertebral line, and three oblong black
spots, margined with white on the sides. Length 18 lines." The same year
DeKa}^ (1842, Part III, p. 70) gives the same characterization. O. P. Hay,
1892, calls its "Form frog-like," and W. P. Hay, (June 20, 1902, p. 128) so
considers the form. To H. Garman (1892, pp. 341, 342) this frog is "small."
This is a rather coarsely built frog, bearing a close resemblance in build to the
Ranidae."

Morse (1902, p. 118) speaks of the crepitans form as "olive brown, with an
inverted 'Y'-shaped green area; the median part of the 'Y' extends along the
vertebral line, the forking taking place on the rump. Brown triangle between
eyes. Sides marked with three oblong blotches. White line from eye to
shoulder . . . Inner surfaces of thighs immaculate. Length i 1/3 inch."

Or M. M. Ellis (1913, p. 58) characterizes it thus: "Head depressed and
pointed, its length about 3 in the head and body; length of the hind leg to
the heel reaching forward to the snout or beyond ; male with a gular sac, size
small, length under 1.5 inches."

COLORATION OF SPIRIT SPECIMENS (191 2)
On the back the color may be grey, light brown, reddish brown, chestnut
or almost black. Rarely green individuals were observed, but in general the
browns predominated. The specimens taken on the open outskirts of the
swamp were greyer while those from the interior of the swamp had shades of
deep brown. All of the specimens over 17 mm. had the prominent black tri-
angular patch between the eyes while almost all of those below 17 mm. to
transformation size had no bars or marks at all on the upper surface which is a
uniform light brown. The triangular interorbital mark has its apex pointed
backward and is bordered by a lighter border which is continued along the

Plate XXIII ,

Cricket-frog (Acris gryllus)

1. Chase Prairie, Okefinokee Swamp, Ga. June i, 1921.

2. 3. Tadpoles, Chesser Id., Ga. June 21, 1922. x i.o.

4. Tadpoles with 4 legs, StarHng Branch, Folkston, Ga. July 24, 1922. x i.o.

5. 6. Transforming and transformed frogs, Chesser Id., Ga. July 8, 1922. Dorsal

aspect. X I.e.

7. Two males and two females, Billy Id., Ga. June 5, 192 1. Ventral aspect, x 0.36.

8. Male croaking, First "Dreen," Chesser Id., Ga. July 25, 1922. Flashlight.

9. Male croaking, Coat-bet pond, Chesser Id., Ga. August 9, 1922. Flashlight.

10. Male croaking, Billy Id., Ga. May 16, 1921. Flashlight.

11. Eggs, Billy Id., Ga. May 16, 1921.

12. Tadpole with two legs, Starling Branch, Folkston, Ga. July 14, 1922. Dorsal

aspect. X I.e.

•d

tf

i^C'

A4^t

Acris gryllus 159

back as a vertical line to the anus. This line is usually brown but may be white
or green. One full grown (No. 6000) example deep reddish brown in color had
no triangular spot, the usual recognition mark of the species. On the upper
lip are three dark bars. Another leads from the eye across the angle of the
mouth to the base of the shoulder. Separated from this by a prominent white
line, another black bar passes over the tympanum from the eye to the shoulder.
Back of the shoulder a dark obhque bar extends two-thirds of the distance to
the hindlimbs. In the adults this mark is likely to remain longer and in young
specimens more likely to appear sooner than the characteristic triangular spot,
at least in the Okefinokee series. Along the back on either border of the
vertebral line are two continuous or interrupted bars which extend almost to
the hind-Hmbs. The hind-limbs are cross-barred, the femur, tibia, tarsus and
foot all having them as in typically colored forms. In the fore-hmbs the
brachium seldom has any bars though the antebrachium may have them. In
the Okefinokee series the most constant color character in transformed in-
dividuals to adults is the coloration of the posterior surface of the thighs.
The typical pattern is upper part with body color with dark cross-bars. Then
follows a series of four parallel longitudinal bands, the first three very promi-
nent, white, brown, white. The fourth brown band may be obscured or absent.
In all the specimens at least one white band shows though the others may be
missing. The under-parts are whitish. Most of the younger individuals are
almost entirely white, but in adults usually the under-surface of the hind legs
is flecked with dark, in a few it becomes very cloudy. Very frequently the
adults have the throat regions also with fine dark dots. Occasionally the
thoracic region and rarely the belly are so marked. The males have yellowish
on the throat. One of the party reported a live specimen with a greenish
white venter.

STRUCTURAL CHARACTERS (WRITTEN IN 19 1 2)

Above, the skin is usually more or less tubercular; some adults have the
dorsal surface smooth as in the young; underneath, gular and thoracic regions
usually smooth, rarely granular or areolate; normally areolation of the belly
is quite distinctly marked off and in some specimens the granulations may form
transverse ridges; thighs are also areolate; nostrils equidistant or slightly
nearer snout than eye; tympanum indistinct; tympanic fold present, fold
across the breast present in about three-fourths of the adults and absent
usually in the young; inner and outer tubercle of foot quite prominent; sub-
articular tubercles moderate ; discs small. The tongue is broad usually notched
behind. In some the notch may be very slight, in others very prominent and
the tongue is then cordate. In one the notch was very broad and was a
shallow depression to each back corner. In several the tongue was round be-
hind or abruptly truncate or obliquely truncate. In one the tongue was
rounded but has a notch at either back corner. The vomerine teeth are
usually two oval patches between or shghtly behind the nares; sometimes the
longitudinal axes are shghtly inclined backward and sometimes not.

i6o Frogs of the Okefinokee Swamp

MEASUREMENTS

(1912-1914)

In 1 9 14 the following measurements were made on the 19 12 and 19 13
material :

"The 44 specimens in length range from 12-24.5 mm., average 20.5 mm.,
mode, 21 mm., the head is 4.75-8.5 mm., average 6.8 mm., mode 7 mm.,
2.5-3.5 in the combined length of head and body; length of head to canthus
oris equal or slightly less than the breadth of the head at the same point,
rarely more than the head; the width of the head 3.75-8 mm., average 6.5
mm., mode 7 mm.; snout 2-4.75 mm., an average 3.8 mm., mode, 4 mm.,
usually 1.5 greater than the diameter of the eye which ranges from 1.25-3.5
mm., average 2.6 mm., mode, 2.0 mm.; interorbital distance 1-2.5 mm.,
average 1.7 mm.; interorbital distance equals eyelid; femur 6-12.75 mm.,
average, 10 mm., mode 11 mm.; tibia 6.5-15.25 mm., average, 1 1.8 mm., mode
12 mm., in every case longer than the femur, usually 1.2 longer; body 1.8
longer than tibia; tarsus 4.5-8.5 mm., average 6.75, mode 7 mm., equal to the
head in length; rest of foot 4.75-15.75 mm., average 9.8 mm., mode, 11 mm.,
longer than tarsus and averages slightly less than one-half the body (.47),
the range however being from .2 5-.9 of the length of the body, comparable to
the two extremes of Cope's analysis ('89, p. 325) for A. g. crepitans and A. g.
gryllus from north and south hut here in one locality. Rest of foot is not longer
than tibia but less: anterior limb 19.5-43.5 mm., average 33.1 mm., averages
1.6 times the length of the body; hind-limb to heel usually extends to or
beyond the end of the muzzle, rarely however, it reaches only to the eye."

(Recent Material)

/fead to angle of mouth 1.3 (2omm.d^) — i.i (20 mm. 9) — 1.07 (24mm.cf)
— 1.05 (24 mm. 9 ) — 1.33 (28 mm. 9 ) in width of head; head to rear of tympa-
num .95-. 85-1. 0-.85-1.0 in width of head; head to angle of mouth 3.2-3.2-2.9-
3.5-3.7 in length of body; head to rear of tympanum 2.3-2.4-2.6-2.8-2.8 in
length of body ; snout . 7-. 7 5-. 74-1 . 1-.8 in first finger ; snout i . 1 4-.85-.98-1 .2-.8
in fourth finger; snout 1.14-1.15-.93-1. 3-1.0 in first toe; eye i. 34-1. 42-1. 8-
1.5-1.8 in snout; eye .54-.70-.66-.83-.71 in tympanum; eye .92-1.07-1.33-
1. 66-1. 4 in first finger; tympanum 4.3-2.8-3.6-3.0-4.0 in intertympanic
width; tympanum 2.5-2.0-2.7-1.8-2.5 in snout; internasal width .66-.66-.86-
.71-.80 in upper eyelid width; interorbital width .83-.71-1.0-1.0-1.0 in upper
eyelid width; interorbital width 1.2 5-1. 7-1. 15- 1.4- 1.2 5 in internasal width;
interorbital width 2.5-2.0-2.7-3.0-5.0 in intertympanic width.

Forelimb. Forelimb 2.0-1. 66-2.0-1. 6-1. 7 7 in length of body; forelimb
3.6-3.0-3.6-3.3-2.9 in hind limb; first finger 2.33-1. 71-1. 75-1. 55-1. 75 in
third finger; second finger 2.0-1. 47-1. 5 2-1. 5 5-1. 7 5 in third finger; second
finger .63-.88-.87-1V0-1.0 in first finger; third finger i. 04-1. 04-1. 02-1. 03-1.0
in second toe; fourth finger i. 4-1. 53-1. 34-1. 4-1. 75 in third finger; fourth
finger .6-.88-.77-.91-1.0 in first finger; internasal width .8-1.0-1. 33-1. 4-1. 4
in first finger; internasal width 1.26-1.13-1.53-1.4-1.4 in second finger; inter-
nasal width 1. 86-1. 73-2. 33-2. 2-2. 8 in third finger; internasal width 1.33-
1.13-1.73-1.50-1.4 in fourth finger.

Acris gryllus i6i

Hindlimh. length 1.8-1.75-1.8-1.56-1.6 in hind hmb; tibia i. 63-1. 63-
1. 66-1. 7-1. 6 in length; tibia 2.8-2.7-2.95-2.8-2.6 in hind limb; tibia .77-
.92-.83-.64-.90 in fore limb; tibia i. 23-1. 4-1. 4-1. 3-1. 25 in hind foot with
tarsus; first toe i. 45-1. 2-1. 44-1. 33-1. 4 in second toe; first toe 2.05-1.74-2.0-
2.1-2.2 in third toe; first toe 3.1-2.6-2.6-2.7-2.72 in fourth toe; first toe 2.0-
1.74-2. 0-2. 1-2. 1 in fifth toe; second toe i. 4-1. 5-1. 4-1. 5-1. 55 in third toe;
second toe 2. 1-2. 2-1. 8-2. 0-1.65 in fourth toe; second toe i. 4-1. 5-1. 4-1. 5-1. 5
in fifth toe; third toe i. 5-1. 5-1. 3-1. 3-1. 24 in fourth toe; third toe .97-1.0-
1.0-1.0-.95 in fifth toe; fourth toe 1.00-.90-1.0-.9 i.o in hind foot; fourth
toe 1. 04-1. 08-1. i-i. 07-1. 28 in tibia; fourth toe .80-1.0-. 92-. 70-1. 15 in fore
limb; fifth toe i. 56-1. 5-1. 3-1. 3-1. 3 in fourth toe; internasal width 1.33-
1. 53-1. 6-1. 7-1. 7 in first toe; internasal width i. 9-1. 8-2. 4-2. 3-2. 5 in second
toe; internasal width 2.73-2.66-3.3-3.5-4.0 in third toe; internasal width 4.1-
4.0-4.33-4.6-4.6 in fourth toe; internasal width 2.66-2.66-3.3-3.5-3.7 in
fifth toe.

HABITAT

Several in their synonymies of this species refer to Wm. Bartram's Travels
(1791, p. 278). Bartram's note follows: "There are yet an extreme diminutive
species of frogs which inhabits the grassy verges of ponds in savannas : these
are called savanna crickets, are of a dark ash or dusky colour, and have a very
picked nose. At the times of very great rains in the autumn, when the
savannas are in a manner inundated, they are to be seen in incredible multi-
tudes clambering up the tall grass, weeds, etc. round the verges of the savannas
bordering on the higher ground, and by an inattentive person might be taken
for spiders or other insects. Their note is very feeble, not unlike the chatter-
ing of young birds, or crickets." This might apply equally to Pseudacris
pcidaris or Bartram's No. 5 frog, "a little grey speckled frog," be Acris as well.

LeConte, its first describer, did not remark in 1825 on its habitat but in
1855 (pp. 426, 427) wrote of A. gryllus that it was "Found in immense numbers
in every piece of water in the Southern States, and was a few years ago com-
monly known by the name of Savannah Cricket."

Harlan (1835, P- 105) says it "Inhabits the Southern and Middle States;
frequents the grass and verges of ponds." Holbrook (1842, Vol. IV, p. 132)
finds "it frequents the borders of stagnant pools, and is often found on the
leaves of aquatic plants, and rarely on the branches of such low shrubs as
overhang or dip in the water." DeKay (1842, Part III, p. 70) has it "fre-
quenting moist wooded places and border of ponds, and is often seen on
aquatic plants .... It is never found on trees, and cannot adhere to the
underside of smooth surfaces."

Abbott (1882, p. 710) contraverts that statement about adherence to
under surface of objects. "In many instances they were found adhering to
the under sides of projecting stones, roots of trees, and even to larger oak
leaves. I find it stated by DeKay in Natural History of New York, that they
cannot retain their hold upon the under sides of projecting objects; that the
discs on their toes are not sufficiently large. This is an error; indeed, the speci-
mens I have in a bottle, can retain their hold when the bottle is turned over."

1 62 Frogs of the Okefinokee Swamp

Cope (1889, p. 329) maintains that "It keeps on the high grass in and
around marshy places, seldom if ever ascending trees or bushes. When pur-
sued it leaps with prodiguous agility, and hides under water."

H. Garman (1892, p. 342) holds "It is more terrestrial than our other
Hylidae, and probably never resorts to shrubs and trees. It is usually found
at the margins of streams or pools, into which it leaps when disturbed, but only
to return to the shore a short distance from the observer."

"In South Florida (Loennberg, 1895 p. 338) along the borders of ponds
and swamps, this frog is abundant though not commonly seen." At Turkey
Lake, Indiana, C. Atkinson (1896, p. 258) finds them "Abundant along the
shallow margins of the lake among marshes and lily pads."

In 1 90 1, Oct. 18, at an artificial lagoon fed by Lake Mendota, Dane
County, Wisconsin, P. H. Dernehl (1902, p. 75) took his series for his Place
Modes of Acris gryllus at Madison, Wisconsin. "Along the banks of this
artificial stream the frogs were extremely numerous, making it a comparatively
small task to secure the one hundred specimens from which my tabulated
results were obtained."

In 1908 Fowler found them in "pools of fresh tide water," in "swamps,"
"about ditches, especially in places where cattle have tramped about the
mud"; also in "dry fields, roadsides and woods."

In Florida near Jacksonville R. F. Deckert (Copeia, 19 13, No. 3, p. 3) finds
this form "plentiful on edges of bayous." In 1915 (pp. 21, 22) he says that
"Acris gryllus LeConte, the 'cricket frog' is one of the commonest frogs, great
swarms of this species having been seen by the writer during the spring months
about the edges of bayous, creeks and ponds."

On Long Island Boyle (1914, No. 7, p. 4) found "Many small ponds vary-
ing from a small pool to a fair sized pond, big and deep enough to swim in. ,
Here as early as March 23, 19 13, I found and collected several "cricket frogs."
Helen Thompson Gaige (1914, No. 11, p. 4) found them "Abundant on the
shores of ponds" in Richland County, Illinois.

In 1917 Hubbs (p. 99) finds them "abundant along the margins of ponds
in the sand dunes between Miller, Indiana and Lake Michigan."

In Texas Strecker (1915, p. 40) finds "This tiny frog is distributed all
over the State wherever there are lakes, ponds, springs or streams. I have
found it even in the heart of well populated cities in little pools formed by
rains." Pope (19 19, p. 97) finds they are fairly abundant all along the banks
and are calling vigorously at Houston, Texas.

In 1923 (p. 36) Viosca states that "Acris gryllus abounds along the creeks"
of his Pine and Hardwood Uplands division or "Shortleaf Pine Hills."

After our 19 12 trip we discussed this phase thus "This species of tree frog
was easily the commonest form of this family in the swamp. It was never
taken in trees or bushes. In the outskirts in the drier lands amongst the
calico-bushes, blue-berries and other heaths it vied with the toads in abun-
dance. Nevertheless, it did not appear wholly terrestrial. On Honey Island,
Floyd's Island and Chase Prairies the full-grown adults as well as transformed

Acris gryllus 163

examples were frequent on the water-leaves along the water-trails we followed.
In the deep cypress forests between Billy's Lake and Minne Lake we noted
them and all over the islands they were very common. Some of our best
collecting courses were the cow and hog trails through the low shrubbery.
Also in the various swampy cross- ways or trails from one island to another
they occurred in the shallow water with the killifishes. Along the swampy
tree and bush borders of Billy Island and in the wooded edges of Lee's clearing
they foraged. In fact, they were as widespread as any Anuran of the swamp
and seemed not to be wholly restricted to land or water. In the main they are
shade lovers except in the open prairies, but here they can rest underneath the
overlapping water-leaves."

In 192 1 our first observation on habitat is April 23. "In the cleaner end
of a cypress pond on the dead black leaves amongst Polygonum, Hydrocotyle,
Pontederia, lots of Ac7'is.'^ Two days later we found Acris common and call-
ing at 10:00 a.m. in a cypress pond (C.A.G. pond. Hyla cinerea, Acris
gryllus, Rana grylio common) . "Took Lucania oryimata in the pond. There are
bonnets (lilies) in the middle, some floating heart, plenty of pickerel weed,
lizard's tail in bloom, water pennywort in bloom, a few southern tickseed in
bloom (Bidens); red bamboo, buttonbush, Virginia willow, etc., around the
edge."

In 192 1 we found them in meadows and around ponds in North Carohna,
in the open vegetation mats or wooded edges of cypress ponds, in moist pine
barrens, occasionally in dry pine barrens, on water prairies, along open lake
edges, on sphagnum strand — in fact, in almost every type of plant habitat
which has any moisture at all.

FIRST APPEARANCE

This topic means very little with this species as the subsequent citations
will show. Whether it be in its southern range or its extreme northern limits
it is very early in the spring or at times appears in warm periods throughout
the winter. Professor T. H. Morgan at Baltimore in 1891 (p. 754) reports
Acris gryllus as the first frog to appear in the spring. O. P. Hay (1892, p.
462) says "this little chatterer appears very early in the spring." In Vigo
County, Indiana, W. S. Blatchley (1891-1892, p. 27) finds that they "are
active in certain localities even in midwinter, lively specimens having been
taken on December 23, January 9 and February 16." In Houston, Texas,
Pope (19 19, p. 97) finds them active Feb. 8, 1918-Feb. 10. "This little frog
was the first to appear in the spring and was abundant along the bayous all
summer,. . ." In Doniphon County, Kansas, Jean M. Linsdale, makes these
interesting observations: "On Feb. 2, 1924, several were found in the creek
above the bridge. They were in the water above some old ice and below a top
layer of new ice. All the frogs were stiff and floating and apparently they were
dead. In the warm water of the springs and just below springs a few frogs of
this species were found throughout the winter."

164 Frogs of the Okefinokee Swamp

GENERAL HABITS

Metachrosis {variations in the individual).

LeConte (1855, p. 427) in describing the extreme variety "in color and in
other marks" of Acris crepitans writes: "Whether all the variations above
mentioned are permanent in the individuals where they have been observed
or depend upon the will of the animal, I have not been able to determine
precisely. In many instances, however, they are the voluntary changes which
for some unknown reason this creature chooses to make in its appearance."

Cope (1889, p. 325) says that "In its habits the Acris grijllus is a lover of
the muddy borders of the water, into which it leaps when alarmed. As it
does not conceal itself among vegetation, like the Hyla pickeringii, it is much
more easily caught than that species, and is more common in museums,
though not less abundant. As the structure of the feet indicate, it is a good
swimmer, and its powers of leaping are remarkable."

Abbott (1882, pp. 709, 710) gave close attention to the subject of their
color and its changeableness. "While there are certain peculiarities of color
that are persistent and characteristic of the species, these become of little
prominence at times, so very great is the difference in the entire coloration of
the animal. Furthermore, they change their hues with great rapidity, and
during the course of a few moments will pass from an ashy paleness or clay
color to an intense black, with the light dorsal stripe scarcely visible, or else
either to a glowing red or brilliant metallic green. So very beautiful are these
changes, and so different will any half dozen prove to be, that it is difficult
to realize that the many before you are one and the same species. Of a series
of six which I have long kept in confinement (October 20th to January 29th)
in a bottle, one specimen was taken from a ledge of pale yellow clay. The
"peeper" was of the same color, the post-orbital dark spot and light dorsal
line being scarcely discernible. The uniform yellow tint, however, was re-
lieved by minute round points of brilliant bronze. This individual, unlike its
companions, did not alter in color for several weeks. The others were very
changeable, and particularly so when exposed to direct sun-light. While I
noted several instances to the contrary my impression is that usually the colors
pale in direct sun-light, and deepen when the animals are in deep shade. This
certainly is true of those I have in confinement, and agrees with my experience
in searching for them during the past autumn. One fact with reference to the
subject of their color is not in accordance, perhaps, with the above, but should
not go unrecorded. The six individuals which I have in a bottle will, at times,
present very different tints, although all are subjected to like surroundings.
Of the six, two or three would be very dark, the others pale yellow. With
some the dark triangular spot between the eyes would be very distinct, in the
case of the others it could not be detected even in outline. It must be remem-
bered, however, that these individuals were kept in most unnatural conditions,
and had, at the time of this writing, been without food for one hundred days
and at the same time remained as active as squirrels.

This species (Cope, 1889, p. 325) possesses the power of metachrosis or
color changes in a high degree. The dorsal stripe and border of the interocular

Acris gryllus 165

spot may be bright green, dirty white, or bright rusty; and the dorsal tubercles
vary in color in the same way. The general tint varies from bright green to
dull slate color. Under A. g. crepitans he remarks (p. 328) "In connection
with metachrosis in this species I observed in a specimen lately dead that on
the end of the muzzle, palpebrae, canthus rostralis, outer Hne of humerus,
ends of sacral dispaphyses, where the derm was in a state of tension, that it
assumed a bright green hue."

Dernehl (1902, p. 81) in writing "Color changes" says "The rapid color
changes attributed to these frogs by various authors as Dr. Hay, Cope and
LeConte were not observed in individuals which we had in confinement in the
laboratory for some time, the only noticeable change being a fading or in-
crease of the ground color, the bands of green or red remaining permanent."

Ditmars (1905, p. 192) notes "This species is subject to rapid and marked
color changes under the influence of varying temperature and the mood of the
individual. These changes affect the general color of the body and the bright
band down the back."

Dickerson (1906, pp. 155, 156) holds that "the dark shades are taken on
when the frogs are in dark situations and especially when the darkness is
combined with a low temperature and moisture; the lighter coloration is
assumed under the influences of bright light, high temperature and dryness."

Variations in color. LeConte (1825) in his original description called its
"colour various." In 1826 he repeats the same.

Almost every author like Boulenger (1882, p. 336) holds its "Coloration
very variable."

Cope (1889, pp. 324-331) gives the upper parts as mainly brown or gray
with diverse dorsal stripes. H. Carman (1892, p. 342) finds the "Size and
color are extremely variable. In most specimens from central and northern
Illinois, the markings are all very obscure, and often the triangular spot be-
tween the eyes is so indistinct as to require close looking to detect it. Others
of the marks described above may even be wanting, and in but few specimens
are all the marks plainly visible. The greenish and reddish forms seem to be
more abundant in southern Illinois. The skin of the more northern individuals
is rougher, the warts often being elongate and ranged so as to form short
ridges."

In another extreme of its range in south Florida, Loennberg (1895, p.
389) reports "All my specimens show the triangular black spot on the head.
The median dorsal stripe is always reddish brown, and I have never seen it
green. The posterior femoral stripes are very conspicuous. The light stripe
from the orbit to the axilla is constant, but the other blotches are not always
light bordered."

In 1902 (p. 128) W. P. Hay calls its "color variable and changeable." So
also Dernehl (1902, p. 80) finds "The color of this species is extremely variable,
usually the upper surface has a ground color of ashy gray with a faint greenish
hue or brown. Dickerson (1906, p. 156) gives green, light red brown, clay
colour, almost black, as dorsal colors.

1 66 Frogs of the Okefinokee Swamp

Ellis (1913, p. 58) has the "color variable and changeable; generally
brownish, greenish or reddish and somewhat clouded with dusky; ..."

On April 23, 192 1, a day or so after my arrival, the following hasty notes
were made on color and environment. "Acris, captured a lot of them. Some-
times on a black soil, and hard to see except when they jump. Some brown
all over back (except for dark marks) when on brown pine needles. Some-
times green all over except for the dark marks. Sometimes gray over drier
sand. Amongst some of the light brown pine needles Acris reddish brown
even on back of forelimbs and hind limbs." A month later on the island May
21, we record "one just captured as black as the blackish earth."

General habits. From C. C.Abbott (1882, pp. 707, 708) we have: "Abundant
as is this batrachian, but little seems to be known of its habits, and certain
misstatements concerning them have long been printed and have never, that
I am aware, been either questioned or contradicted."

"During the month of April, 1881, 1 had excellent opportunities for observ-
ing these little creatures, and finding that but httle had been recorded concern-
ing them, availed myself of my chance and watched them closely for several
weeks."

"While a network of ditches in a low meadow were being repaired or

cleaned, I followed the workmen closely, the one striking feature

of the locality, at this time, was the wonderful abundance of little "rattlers"
(Acris crepitans) , as I prefer to call them .... From the date of their earhest
appearance until May 20, their numbers were incalculable. In every portion
of the meadows at all wet, they were to be seen. Extremely active and very
shy, they were difficult to catch provided you pursued a single individual,
but by sweeping an ordinary dip net along the grass at the edge of any little
pool, several were certainly to be caught .... About the 20th of May there
was a very noticeable dimunition of their numbers, and by the loth of June
not a specimen was to be found.

"The fact is, that their vigor culminates with the maturity of the ova and
spermatozoa, and having spawned, they have no vital force remaining, and in
the course of a few days after ovipositing, they die. Weeks then elapse when
no representatives of this batrachian are to be found; indeed none exist,
except the thousands of tadpoles.

". . . Their physiological activity culminates with the maturity of the
ova and the labor of depositing it; this effected, they are worn out and in a
very short time, die."

The article of which the above are excerpts is very interesting and in
general instructive, but we believe this author falls in error in beUeving the
adults die after breeding. Sometimes frogs of most species are found on
breeding grounds dead. This is from vigor of breeding; such as violent
matings, individual exhaustion after breeding etc., but a general rule of
death does riot obtain for any Anuran of Canada or the United States. We
have captured Acris adults every month of the year and spent females
aplenty. If there be any lessening in numbers it may be because of migra-
tions, vegetation more concealing and the enemies of which he speaks.

.4cm gryllus 167

O. P. Hay (1892, p. 462) finds: "This little frog is one of our commonest
batrachians. During the summer season it may be seen in numbers along all
of our streams. I doubt if it is often seen about the ponds or pools far from
running water. It is not thoroughly aquatic but delights to spend its time
amid the vegetation about the borders of the water. When alarmed it will
leap into the water, but it often appears to become alarmed at its rashness and
hastens to reach the land again. When followed up, however, it will go to the
bottom and seek to conceal itself for awhile. Though belonging to the
'tree-frogs' it never ascends trees, and probably climbs only the shorter
grasses and water plants." Blatchley (1891, p. 27) calls this species "the
most abundant tailless batrachian in the country. Hundreds are to be seen
along any small stream in spring and autumn. They appear less common
in summer . . ."

"... It is my observation that it is to be found at all times during
the summer."

Hurter (1893, p. 253) finds this form "the most common of all little
frogs. It is found near pools and creeks, where it generally sits on the banks
so as to resort to the water when alarmed. It can make jumps from three to
four feet long."

Sherwood (1898, p. 18) holds "They are rarely found away from the bor-
ders of ponds, and their long leaps and swimming powers render capture very
difficult."

Morse (1902, p. 118) finds that "So nearly does the color of the frog blend
with that of its surroundings that it is detected with but the greatest dif-
ficulty. In the cricket frog we have virtually a tree-frog with terrestrial
habits. The presence of the terminal suckers on the toes would seem to indi-
cate a former arboreal mode of life. Common everywhere."

Ditmars (1905, p. 193) says "The animal is very difficult of capture since
it possesses great leaping powers and quickness in diving. During periods of
heavy dew these frogs may be found in high grass adjacent to marshes. Owing
to its smallness and agility, it is rarely observed after its breeding season."

Of this species Fowler (1907, p. 102) writes "These little toads were found
to be very abundant, though exceedingly difficult to see, on account of their
small size and close resemblance to the aquatic vegetation in which they live.
Their notes were a characteristic feature of these ponds all along the marshes
most of the time, and also in the cedar-swamps. I have heard them in
various places, more or less regularly, from spring all through the summer
and on several occasions in Dr. Abbott's own meadows." (See Abbott 1882).

In 191 8 Cope (19 19, p. 97) writes of them at Houston, Texas, that "The
frogs usually sit on the bank a few inches from the water. If danger ap-
proaches they plunge into the water and swim rapidly back to the bank unless
too much alarmed, when they dive and hide in the mud. February 10 col-
lected seven specimens in day fight. They are not nocturnal in habit but seem
to be active both day and night."

Of the jumping records of Acris we record the following: On Apr. 23, 192 1,
we wrote "Acris can jump three or four feet sometines. . . . Acris usually

1 68 Frogs of the Okefinokee Swamp

jumps for several leaps before it disappears." On Apr. 25, 192 1, we repeat
that Acris males can jump 3 feet at times on water's surface." On May 17,
we compared Acris's leap with our longest stride and wrote "Acris can leap
at least my own pace." At another time some adult Acris puzzled me because
they leaped into bushes 1-2 to i foot high." We queried as to the possibility
of their being Acris one-year-old or Pseudacris but they were neither. On
May 21, 192 1, we note "Acris has been on the island hopping around on the
ground and into small bushes from the ground and down to ground again.
May be some of these have left the ponds."

Most writers assert that they never climb. On June 28, 1922, at Coat Bet
Pond on Chesser Island, where flags were reaching above the water we found
six or eight Acris crawled up on flag stems 2-3 inches above the water's sur-
face. Sometimes they would leap off. Usually they stayed in place.

VOICE

Holbrook (1842, Vol. IV, p. 132) writes that "This is a merry Uttle frog,
constantly chirping hke a cricket, even in confinement; . . ." Of some
captives he says "Their chirp, at times, was incessant, and sprinkling them
with water never failed to render them more lively and noisy." The last
two expressions of "lively and noisy" DeKay repeats.

Charles C. Abbott (1882, p. 707) remarks that "One of the earliest indi-
cations of returning spring is the dear, bell-like note of the little batrachian,
called by many the "Savannah cricket," known in New Jeresy as the "peeper"
and scientifically designated Acris crepitans Baird." . . . "They were in
full song, (April 1881) and when not disturbed, made more noise than all
the frogs in the neighborhood together. They are quite timid, however,
and on being approached were straight way 'mum.' Their vocal efforts
seemed to increase until about May ist . . .."

Of .4cm gryllus crepitans Cope (1889, pp. 328, 329) notes that "The
note of this species may be exactly imitated by striking two marbles together
first slowly, than faster and faster, for a succession of about twenty or thirty
beats. The noise can not be heard at a very great distance. Like Hyla
pickeringii this species in confinement can readily be made to produce the
note by imitating it, either with the voice or the clattering of two pebbles."

Hay (1892, p. 462) says "It is a cheerful little creature, and on warm days
may constantly be heard executing its noisy song. This resembles closely the
striking together rapidly of two pebbles, and, often, when their singing has
been interrupted by the passer-by, it may be started again by clicking two
stones sharply together."

H. Garman (1892, p. 342) writes of voice as follows: Its note is a rapidly
repeated grating noise, thought to resemble the trilling of a cricket, whence
the name cricket-frog."

Sherwood (1898, p. 18) writes "Very early in the spring and before the
appearance of the 'peepers' {Hyla pickeringii), their rattling, broken cry may
be hard. It is not sharp like that of Hyla, and would not be noticed unless
one were near."

Acris gryllus 169

W. P. Hay (1902, p. 128) writes that "Both (subspecies) are found in
abundance along the banks of streams and ponds whence their continuous
metalHc cry comes almost without intermission during the season of activity,
particularly during hot, sunshiny days."

Moi'se (1904, p. 118) remarks that "The cricket-frog is well known to any
one who frequents the riverside or the swamp — less, however, by its appear-
ance than its note. If one can imagine a rattling of pebbles mingled with the
screech of a violin string in a high note, he may have a suggestion of the
cricket-frog's note. When given it is either a continuous chirp or given in
sets of three each rising in pitch."

In 1905 Ditmars speaks of "its sharp, trilling cry, resembling in volume the
call of the field cricket, ... It "may be heard at various times."

Ruthven, Thompson & Thompson (191 2, pp. 46, 47) report that "These
frogs sing in chorus during the months of April and May and the isolated call
may be heard all summer. The song resembles the chirping of a cricket,
hence the common name, cricket frog." To Miss Dickerson (1906) the song
"resembles the ratthng call of the Swamp Tree Frog, but the notes are more
rapidly given and are sharper in quality." She gives the pebble or marble
figure. "The call has not great carrying power, either when given alone or
when given in chorus."

Fowler (1907, pp. 102, 103) finds "Besides the rattling call a squeaky sound
was heard occasionally, though only during the breeding season. ... It
may be stated that in my experience their call appears to be variable. . . .
(two pebbles figure). . . . Perhaps the rattling of two castenets would be
a better suggestion." Deckert (1915, p. 21) finds "Its rattling notes can be
heard during the entire season, day and night" in Florida.

At Brevard, N. C, Dunn (191 7, p. 621) from July i "Heard (them)
nearly every night."

In 1920 Logier (1925, p. 92) found one specimen of this species at Point
Pelee, Ontario. "On the afternoon of July 15th, while sketching beside this
pond, my attention was caught by what to me was a new frog-note, and which
sounded something like the tapping together of two pebbles. This call,
which like the calls of our other diminutive frogs had considerable carrying
power, was uttered intermittently, that is, the frog would call several times
and then remain silent for a while. By following the sound I finally located
the tiny producer, hiding among the rushes in the shallow water at the pond's
margin. It dived when I attempted to capture it, but was dragged ashore in a
net full of bottom trash. When landed it did not make for the water as frogs
usually do, but took a few jumps further ashore as if in an effort to hide itself
in among the shore vegetation."

Our records of calling in 192 1 and 1922 extends from Apr. 21 into Septem-
ber or the whole period of our sojourn in the swamp.

Characterizations of its voice are quite numerous. For example, Over-
ton's description is one of the most extended and best accounts: "The voice
of a cricket frog is a combination of a rattle and a musical clink, but it is only
about half as loud as that of a spring peeper. A chorus heard at a distance

lyo Frogs of the Okefinokee Swamp

sounds like the jingling of small sleigh bells, for the musical element of its
call travels farther than the rattle. A chorus heard close by sounds like the
rattle of small pebbles poured upon a cement pavement.

"An individual frog sings for from thirty to forty-five seconds at a time.
Its call has three phases. The first phase lasts for about five seconds and
sounds like the clicks of a boy's marble dropped upon a cement pavement
once or twice a second from a height of about six inches. The second phase
sounds like the galloping of a small pony on a brick pavement, or like the
clicks of a boy's marble dropped upon a pavement from a height of only an
inch or two, and allowed to bounce twice each time. The third phase sounds
like the regular cree-cree-creeing of a tree cricket, or like the rattle of a boy's
marble that bounces rapidly when it is dropped at frequent intervals from a
height of only half an inch. The time and rhythm of the sounds are about
the same as that of the following syllables pronounced with the speed of
ordinary reading; — "click, click, chck, click, . . . click-e-ty, cHck-e-ty,
click-e-ty, click-e-ty, click-e-ty, . . . cree, cree, cree, cree . . ."

"The cricket frog inflates a vocal sac under its chin during its call. It often
sits quitely with its sac distended for many minutes between its calls. The
violent efforts of its body in producing its sound make the frog resemble
a small boy on his hands and knees blowing a fire with all his might. The vocal
sac is bright yellow and when it is seen distended in the day time, it is so con-
spicuous that it reveals many a singer that otherwise would be almost in-
visible on a lily pad."

To us, on April i6, 192 1, at Raleigh, N. C. "Acris gryllus calls sound like a
rattle or some of the metal clickers." On April 25, 1921, in the Okefinokee
Swamp, we repeat "Acris call is a rattle," just as C. M. Barber, 1923, speaks
of the "rattle of Acris." On May 18 I have the following journal notes:
"Acris call. Tick, tick, tick, tick and so on. Or kick, kick, kick, kick, . . .
kick, kick . . . kick, kick, . . . kick, kick, . . . kick, kick, kick, kick,
kick, kick. One Acris went on calling for 45 seconds. Another kick, kick,
kick, kick, . . . kick, kick, . . . kick, kick, kick. Another 4 kicks pause,
then 7 kicks in succession. Another called for 30 seconds straight." To F.
Harper "its ordinary note is a simple rather strident gik, given either singly
or in irregular but successive series.

Frequently one finds the males with inflated vocal sacs even when not
calHng. When calling the throat is never fully deflated. After a call it may
be swollen to three-quarters the full capacity. Thus when the call is given
the body sides are compressed and the vocal sac extended to its limit.

Our voice records for 192 1 and 1922 follow:

Apr. 15

" 17

" 21

" 23

" 24

" 2 s

I92I

Raleigh, N. C. Males calling.

Raleigh, N. C. Plenty calling in evening at Boone's Pond.

Billy's Island, Okefinokee Swamp, Ga., Voice 5:00-12:00 p.m.

Last night in early evening went after croaking Acris.

Moonshine Pond. In Pond heard no end of Acris 7 :3o p.m.

In Long Pond at 10 : 00 a.m. heard quite a few Acns caUingin Hylacinerea

pond. In C. A. G. {cinerea, Acris, grylio) pond Acris common and calling.

Acris gryllus

Apr.

May

day temperature 66 -69 .

May

June

July

Calling at Billy's Lake.

Some Acris heard around Long Pond.

One heard.

7:00-8:00 P.M. Air 58°, few .4cm heard.

Several heard at Billy's Lake.

Several heard.

Acris in strong force 6:45 a.m. Air 64°.

This evening at 8:00 p.m., 72°, no end of Acris calling.

Swimming pool. Plenty of calling Acris. Afternoon.

Acris commonly calling at swimming hole and mating.

Temperature tonight only 66° and 67.° Only ilcns calling. During

C.A.G. Pond.

:30 several heard.

[n strong chorus
Heard on the lake.

Few Acris calling occasionally in the ponds.
Minne Lake at noon. Several calling.
Acris still going. Air 77°F.
No end of Acris calling.
Common and in good voice.
Heard a few tonight.
Acris — several heard.
Acris common along canal.
One calling.
Plenty of Acris calling.
Mixon's Hammock. Hear Acris.

Minne Lake 10:55 a.m. hear several. Many in afternoon.
Several on prairie (Floyd's).
Floyd's Prairie 7:55 a.m. Several heard.
Several calling at lake.
Long Pond some calling.
One heard.

Plenty calling aloud Chorophilus Pond.
One heard. A few occasionally calling.
One heard.
A few heard.
One heard.

An occasional Acris gryllus at Long Pond.
Several heard.
Two heard.

Last night great chorus.
Abundantly heard on trip to Chesser Island.
Acris common Chesser Island.
Acris abundant in cypress pond.
Acris very active.

Acris calling as lustily as Hyla femoralis.
Many heard.

One heard at noon (rain).
Last night quite a few Acris.
Acris commonly heard on prairie this afternoon.
Acris at Black Jack Island.
Several Acris heard Black Jack Island.
Acris calling.
>3. Heard down Suwannee River and along St. Mary's River from source
to mouth by F. Harper and Marion Lee.
Aug. 23 . Acris only anuran voice tonight.

Aug.

172 Frogs of the Okefinokee Swamp

1922

June 13 . Acris heard on Chesser Island.

" 14. Common at Coat Bet Pond.

" 22. In great chorus in Coat Bet Pond and on the prairies.

" 27. Chorus of them tonight.

July I . One heard in cypress bay.

" 2. Many heard calling and mating at Starling Branch.

" 3 . Commonly heard.

" 4. Several heard.

" 5. Abundantly heard.

" 6. Acris frequently heard on prairie and about "houses."

" 7 . Grand Prairie — many calling.

" 9. Common, active at Coat Bet Pond.

" II . Several heard.

" 12. Calling moderately at Coat Bet Pond.

" 13 . One heard.

" 18. In chorus at Coat Bet Pond.

" 19. Spanish Creek — several heard.

" 21 . Several heard near Dinsmore, Florida. Abundant at Coat Bet Pond.

" 22. Several heard at noon.

" 23 . Few calling early.

" 25. Some calling after rain.

" 26. Several calling at Starling Branch.

" 27. Several calling near dark.

" 28. One or two calling.

" 31. One or two calling.

Aug. I . Calling commonly along Dixie Highway near Waycross.

" 2 . Calling on Billy's Island.

" 3 . Several calling in grassy field.

" 4. Calling in considerable numbers 11:15 P-^^-

" 5. 10:00 p.m. — Many calling.

" 6. Common on Billy's Lake.

" 7 . One calling.

" 8 . After today's rain all along Okefinokee Road Acris calling 8 : 00 — 1 1 : 00
p.m.

" 9. Trader's Hill Pond Acris calling 3:15 p.m. Calling lustily 8:00-11:00
p.m.

" 10. Abundantly calling Petty Pond 8:45-10:30 p.m.

" II. Common from Folkston to Chesser Island but not at their height.

" 12. Several calling during day.

" 13. Commonly heard from St. Mary's, Ga., -Chesser Island 7:00-10 :30 p.m.

" 14. Several at night.

" 15. Common near Callahan, Florida.

" 18. Heard in pine barrens.

" 19. Commonly heard near Lake Sego.

" 21 . One heard.

" 23 . One heard.

" 28. Acris calling at Coat Bet Pond 9:00 a.m.

" 29. An occasional Acris.

" 30. Acris common on prairie and islet, commonly heard at night.

Sept. I . Heard at night.

" 2 . Several heard at night.

" 3 . One or two heard on islet and others on prairie.

" 4. Two on camp on islet.

Acris gryllus 173

Considering 16 choruses from April 24- July 18, 192 1, we have for nearby
station records of maxima 8i°-95° with exception of two 73°, usual range
8i°-88°, average 81°, mode 86°, for the day before; and minima from 5o°-73°
(one to 46°), average 60°; for day of record, maxima of 81 "-95°, average
89°, minima of So°-73°, average 69°. Heavy rains preceded about half of
the choruses. In last of April temperatures 84°-87° maxima and 5o°-62°
minima obtained the day of the records. In the first half of May maxima
8i°-9o°, minima 53°-67° prevailed. In the choruses of May 26 and 27
maxima 93°-9S°, minima 65°-7o° prevailed. In July 11-18 maxima 87^-94°
obtained, minima 68°-73°. Minima thus mount from 5o°-62°, 53°-67°,
65°-7o°, 68°-73° in mid July. The mid July choruses were started by rains
of two inches or more.

The actual records of air temperatures taken by us in 192 1 when we heard
Acris were about 10 instances from April 30- July 29. They range thus:
58°, 72°, 66° or 67°, 71°, 80°, 74°, 70°, 74°, 76°, 80°, 73°. In June and July
each of these were preceded or accompanied by rain, clouds, storms, or
threat of lowery weather. These average about 72°.

In 1922 on Chesser Island from June 14-22 we heard great choruses on
the island and on the prairies and later in the later part of June. No minimum
during our waking hours was below 7i°-73°. Many days went up to 90° or
higher, and much rain or cloudy weather was on. It seems that in mid summer
rain is a potent factor. They will peep at 56° or lower. Somehow 65°-7o°
seems to be a preferable temperature for cheerful Acris. When minima get
to 65° or 70° or higher and maxima reach 80° or 85° on into the 90^'s with
some rain we have optimum conditions for great choruses.

MATING

Male {From life, June 25, 1921). Stripe down back and around triangle
dark olive buff, throat raw sienna. All over the throat collections of dark
dots which sometimes arrange in a reticulate fashion. Iris pale vinaceous
drab on black. Light orange yellow rim around the pupil.

On one day, Apr. 23, I saw black, dark brown, reddish brown, light brown,
green and gray specimens of Acris.

Female {From life, June 25, 1921). Of a mated pair. Clove brown above.
Triangular spot between eye obscured by this dark color. Throat, breast and
belly pale olive buff. More or less on under side of fore legs same color and so
the spot below eye and along the upper and lower jaws, also the line from
eye to arm insertion. Area back of arm insertion palHd vinaceous drab.
Oblique bar on side clove brown with pale olive buff and oHve ocher: Under
side of hind legs clear with a Httle pigment. Brown long stripe on rear of
femur snuff brown or dresden brown with clear unpigmented stripe below and
above. Also another such brown stripe above the upper clear area. From
vent around bases of hind legs to venter are pale oHve buff papillae.

Structural differences. H. Garman (1892, p. 341) is one of the first to note
that "The first finger of the male (is) but slightly swollen."

174 Frogs of the Okefinokee Swamp

Many remark the difference in the throat of the two sexes. ElHs (19 13,
p. 58) holds "its voice being powerful because of the gular sac."

On Apr. 25, 192 1, a pair were captured. Of it we record that "At night
the Ac7Hs look all faded out including femoral stripes." "Male, orange
yellow throat with white and dusky flecks. Forelimbs of male look slightly
larger proportionally than those in the female. Often male may be green on
back except for cords on back." On May 15, 192 1 we write "The Acris males
have the orange throat; this orange color extends to the brachium. Some-
times the skin on the brachium is papillose or raised up into little processes."

The measurements of loi adults from Okefinokee Swamp, i.e., 55 males
and 46 females yield the following results: males range from 15-24.5 mm.,
mode 18 mm., average 18 mm. ; females, range from 15-24 mm., mode 21 mm. ;
average 19 mm. A study of the modes reveal that the males are at 18 mm.
and the females at 21 mm. There are four doubtful external determinations
of males from 15-17.5 mm. and of females four such determinations. At
15 mm., however, there are two certain males and one female; at 16 mm. 10
certain males and no females, at 17 mm., 8 certain males and 6 positive
females. The range of 15-17 mm. is the uncertain overlap of transformation
size with the beginning of maturing breeding frogs.

In a group of adult Acris collected from New Orleans by Dr. R. W.
Shufeldt (U. S. N. Museum No. 13270) the 22 males ranged from 19-22 mm.,
the modes 20 and 22 mm., average 21 mm., the 30 females were from 16-26
mm., the modes 22 and 23 mm., average 22 mm. The modes and averages for
males and females were higher than those of Okefinokee.

A collection made at Ames, Iowa, June 12, 1924, by Drs. E. L. and K. V.
(Katherine VanAVinkle) Palmer give a higher modal and average size. The
twenty-two adults were taken from a breeding congress, 18 males and 4
females. The males range from 21-29 mm., mode 25 mm., average 25 mm.,
and the four females ranges from 27-29 mm., average 28 mm.

The measurements for 169 adults from various parts of Acris' s range,
i.e., 100 females and 69 males gives: females range from 16-33 mm., modes
22, 24, and 28 mm. average, 23.5 mm.; males range from 17-29 mm., modes
20, 22 and 25 mm., average, 22 mm.

Compared to this composite series the Okefinokee males have their range
and mode 2 mm. lower, and their average 4 mm. lower; the females their
range and mode i mm. lower and their average 2.25 lower.

Many queries arise: Do the 13.5, 14, 14.5, 15 mm. transforming individuals
becomes sexually mature at once or are the debatable and positive 1 5-1 7.5 mm.
cTs from frogs which transformed at 9-12 mm.? Are all the adults from 15-
24.5 mm. (the largest) one-year-olds when breeding or is less than a year
needed for maturity?

At 15 mm. some males and females are not determinable externally.
An analysis of a group of 30 frogs we took from Apr. 25-May 3 follows. These
sex identifications are on external characters alone.

13 mm. not at transformation; 15 mm., possibly male; 15.5 mm., is it a
female? 15.5 mm., is it a male? 16 mm. male two central plaits, throat yel-

Acris gryllus 175

lowish; 16 mm., sex not determined. Is it a female? 16 mm. looks a gravid
female; 16 mm. probably male, heavily spotted from throat to fore abdomen
but no plaits and little yellow on throat, yellow under hind legs, 16.5 mm.
Is it a male or female? 17 mm. Is it male? Yellowish under hind legs; 17
mm. looks like female; 17 mm. looks like female; 17.5 mm. female, gravid,
spotted throat and breast; 17.5 mm. sex not clear; 18 mm. male, throat
heavily spotted, a single central plait; 18 mm. looks like female; 18 mm.
female, spotted throat and fore abdomen; 18 mm. male, yellow throat and
under hind legs; 18 mm. male, yellowish on throat, two lateral throat plaits,
heavily spotted on throat, lightly so on abdomen, thumb possibly larger,
under side of hind legs yellowish; 18.5 female, faintly specked on throat and
pectoral region. A very rough individual; 19 mm. male some color on venter
and throat, a median plait and a lateral plait on either side of throat, heavy
spotting on chin and fore abdomen; 19 mm. female almost without specks
on under parts; 20 mm. female, throat feebly punctate; 21 mm. female throat
and pectoral region spotted; 21 mm. female throat and fore abdomen faintly
spotted; 22 mm. female throat to fore abdomen faintly specked; 23 mm.
female throat like rest of underparts; 23 mm. female throat, pectoral and fore
abdomen heavily spotted.

The plaits on the throat may be one median plait or as often two central
plaits. Occasionally beside one or both of these two will be a subsidiary one
parallel with it. Sometimes under one ramus will be a parallel plait or one to
each ramus. The throat may be very yellowish bulfy. In alcohol it looks
quite dark at times or may appear smoky or grayish with clear white papillate
dots and fine punctures.

Some of the measurements of mated pairs are: 20.5 mm.d^ X 24 mm. 9 ;
18 mm. cf X 22.5 mm. 9 ; 19 mm.cf X 19 mm. 9 ; 18.5 mm. (^ X 21.5
mm. 9 ; 20 mm. cf X 24 mm. 9 .

Duration, night or day. On the evening of Apr. 24, 192 1, we went to
Moonshine Pond "Where we heard Hyla femoralis in the trees. In the pond
we heard one Hyla cinerea calling. No end of Acris calling. Some Ra7ia
sphenocephala and plenty of Bufo lentiginosus began calling later. This at
7:30 P. M. Later captured on lily pads 5 calHng males of Hyla cinerea. In
same place took my first Acris pair in Okefinokee Swamp but they got away
from me. In the pond were plenty of water lilies and water pennywort
(Hydrocotyle)."

On the following day at 8:00 p. m. and later in another pond "on the lily
pads and vegetation-carpet were untold numbers of Acris gryllus. The male
often would keep the sac much inflated even though not croaking. Finally
found a pair in axillary embrace. These were put into a bottle but they
broke. At camp they were mated."

On May 14 in another pond at 8 : 00 p. m. "The Acris were calUng loudly.
We found among lizard's tail {Saururus cernuus) on the sphagnum surface-
mat, a pair. They were resting quietly. Put them in a butter jar. The top
of the jar had a small hole about which we had forgotten. When next we
looked the pair had escaped. The next pair we saw amongst iris or sphagnum

176 Frogs of the Okefinokee Swamp

at the surface. Tried to catch them with both hands and failed. Later
found another pair; when the electric flashlight went out they leaped away.
They can leap two feet or more when mated. Finally caught the pair. Put
them in a sack. This in a jar with hole plugged .... We took the pair of
Acris out of butter jar and put them in a porcelain jar for two flashlight
photographs. Later transferred them to photo jar without their breaking
embrace."

On May 23, 192 1, at a bayou of Billy's Lake we observed the odd behavior
of two croaking males. Our journal note with no poHshing is thus: "At
noon along the edge under shade of some bushes was a small series of grass
blades 5 inches high. A pair of Acris males, one directly behind the other
about 3-2 inches. Each croaking male revealed his yellow throat very
plainly in day time. Then it leaped on to the other. Then it leaped to one
side and they were more or less back to back. At first I thought a mated pair
must be in the water back of them because of water commotion, but it was
each male stretching one leg out as far as he could behind. Then the
second male attempted to embrace the other but merely leaped on to it.
This process kept up for the 20 minutes we remained. Once one male crawled
up the grass. The other followed. Once on a leaf-blade one could fairly see
the leg stretched back. So also at other times. At times the leg would remain
thus stretched out for some time, "frozen" so to speak. When another male
not far away began croaking both croaked at same time. The process of play,
by stretching, leaping on to each other or about each other was common to
both of them."

On May 26, 192 1, with air temperature 94° in partial shade 4:00 p. m.
we found "gravid females in the pine barrens. About the ponds were no end of
Acris calling. Saw two mated pairs. One at the base of a black gum on wet
dead leaves at the edge of the pond and one pair on top of a grassy mat
where many males were. Every mat had a male or two perched on it. The
pair captured broke. At midnight they were not remated." On the next
day. May 27, we have "The Acris pair of last night apparently have resumed
mating since I caught them. The female has laid a few eggs."

At Southern Bullfrog Pond on the evening of June 3 "we found two mated
pairs of Acris. Lost one. At Chorophilus Pond, the same evening were
mainly Acris. Captured a pair there."

In 1922 we found a mated pair July 2 at Starling Branch crossing, between
8 : 30 p. m. and 1 2 midnight. "In a grassy overflow with Bufo terrestris and Hyla
femoralis were plenty of Acris. Finally at 10:00 p. m. found a pair at the
edge of the pond. Photographed them. Brought the pair to camp in a small
bottle. They were caught in the bottle. The female and male broke. Stop-
per made it too confmed and the female died."

Our experience at Dinwiddle, Va., June i, 191 7, may be apropos. "Toads,
tree toads and bullfrogs around. ... At the first ford about 6 miles be-
yond Dinwiddle, Va., near the road found several files of Fowler toad's eggs.
. . . Found a mass of bullfrog eggs. . . . (tree toad eggs in another pond) .

Acris gryllus 177

"After supper Mimz (Professor P. A. Munz, Pomona College) and I went
out for frogs. ... In the Acris pool, long shallow, 1-4 inches deep
(J uncus plentiful) we found plenty of Acris. After about one hour's collecting
we found a mated pair in the water. The embrace is axillary .... Didn't
find any of the eggs. The pond is a surface pond with trees on all sides but
the east. . . . About 11:00 p. m. returned. Put extra males and female
Acris in one compartment and started to put the pair in another part when we
lost the female of the pair. A severe storm came up. Later Munz and I went
out again. Soon found another pair. After capture they broke but soon
resumed. When we put them in the can they separated. In the morning ( June
2) no eggs from Acris. Munz and I went to the Acris pond and found the
eggs." (See egg mass description). The mated pair never laid but three days
after, June 4, 191 7, one of the females laid eggs. These were our first identi-
fications of Acris eggs in the field.

Amplexation. Normally Acris embraces are as in the Hylidae in general,
i.e., axillary amplexation. The Acris pairs taken at Dinwiddle, Va., in 191 7
were all axillary in type. On Apr. 25, 192 1, a pair captured on a lily pad "was
found in axillary fashion. When I put them into a bottle they broke. At
camp they were mated axillary fashion. Then put them into a compartment
of the fish can. The male then held the female with his forelimbs ahead of
the forelimbs of the female and the hand of each forefimb came on to the
pectoral region." The amplexation of all four pairs of Acris found the
evening of May 14, 192 1, were axillary with no departures. So also were the
two pairs of May 26, the two pairs of June 3, and the pair of June 25, 192 1.

OVULATION

Habitat. Abbott suggests blades of coarse grass. This expands in Sher-
wood to grass and weeds and in Ditmars to grass and reeds. John K. Strecker
Jr., (19 10, p. 80) places the "breeding locahties" as "ponds and small streams."
In 1906 Miss Dickerson (p. 156) has the eggs "attached to grass blades or
leaves in the water."

In 192 1 we took mated pairs in the open area of cypress ponds, in amongst
the cypress of the edge of cypress ponds and bays, in cypress bays or glades,
on sphagnum strands on prairies, in wooded streams or branches, and rarely
in moist pine barrens. The open cypress ponds and prairies were the optimum
habitats for ovulation.

Period. Abbott (1882, p. 707) gave this period as about May i. Hay
writes "Numerous specimens were found at Irvington on the 8th of March.
The eggs are probably laid about this time, although I do not know anything
about them." Brimley (1896, p. 501) says "This species (in North Carohna)
breeds from April through most of the summer." Doubtless W. L. Sherwood
(1898, p. 19) is following Abbott when he says "The eggs are laid early in May,
in small bunches attached to grass or weeds. Development is prolonged."
Following Abbott and Sherwood, Ditmars (1905, p. 193) writes "The eggs
are deposited early in May in small bunches. They usually adhere to grass
and reeds."

lyS Frogs of the Okefinokee Swamp

In 1906 Dickerson paraphrases the above data of period and asserts egg
laying is in late April and early May.

In 1910 J. K. Strecker Jr., (1910, p. 80) gives "Apr. 10 to May 30" as the
breeding period for Aais grijlhis crepitans of Waco, Texas. Ruthven, Thomp-
son and Thompson (1912, p. 47) hold "the eggs are laid in April and May."

In 1920 Wright (1920, p. 29) writes "The cricket frog is one of the first
forms to appear in the spring, and according to several authors breeds in
March, April and May or even later."

In 192 1 on April 15, on St. Augustine School Grounds, Raleigh, N. C,
we heard a few males calling at night 9 : 00 p. m. with temperature 68°. Then
we thought that they and the meadow frogs were practically over breeding
with a few stragglers in the ponds and wet meadows. But two nights later
at Boone's Pond plenty were calling in the evening. This indicates Brimley's
remark that they "breed from April through most of the summer" is nearer
the truth for Raleigh.

In 1 92 1 in the swamp from captive pairs we have May 15, May 16, May
27, June 4, June 25. In 1921 we took or recorded mated pairs: Apr. 24,
Apr. 25, May 14, May 15, May 26, June 3, June 24. In 1922 we captured a
pair July 2.

In 192 1 we recorded great choruses Apr. 24, Apr. 25, May 9 and 10, May
13 and 14, May 16, May 26, May 27, June 3, June 25, July 11-18, July 27.
In 1922 the choruses were June 14-22, June 27, July 2, July 5, July 7, July 9,
July 18, July 21, Aug. 4-6, Aug. 9-1 1, Aug. 13, Aug. 15, Aug. 19, Aug. 30.

On the basis of spent females we find that in a group of 9 females captured
Apr. 24-28, 192 1, the night after we entered the swamp, 7 were gravid and 2
spent. Other female records for 192 1 are June 16, gravid, July 2, gravid,
July 29, gravid, June 3, spent, July 17, gravid, June 20, gravid, July 8, gravid.

We, therefore, see that on the basis of ovulation in camp we have records
from May 15- June 25, on mated pairs evidence records from Apr. 24- July 2,
on choruses from Apr. 17-Aug. 30, and on spent and gravid females records of
spent females Apr. 24 and gravid females July 29. The ovulation period
surely goes from Apr. 2 4- July 29 and if strong choruses help in the evidence
from Apr. 15-Sept. i. All our subsequent notes (1922-1926) summarize
it as Apr. 15 or earlier to Sept. i. Doubtless it lays beyond Sept. i.

Temperature and humidity. In 192 1, May 15, when the first pair laid in
captivity the air temperature was 67°. Later the same day when another
female laid we report "Today the temperature has been from 66°-69°. Still
later, May 16, we record "The eggs during May 15 and May 16 to 9:00 a. m.
had air conditions of 67°-7i°F.

Humidity is the most important factor. The Apr. 24-25, 192 1, period was
preceded by heavy rain in surrounding stations. The May 13-16 was rainy
and was followed by rain. The May 26-27 P^^riod had a few light rains
around the swamp. On June 3-4 we had heavy rains. On June 24-25 rains
prevailed. In 1922 on July 2 we had a rain at St. George of 2.00 inches.
The air maxima and minima for stations around the swamp for these peri-
ods are :

Acris gryllus

179

Air Mi

ixima

Air

Minima

Range

Average

1921

Range

Average

81-84

84

Apr. 24-25

50-62

54

73-88

82

May 13-16

59-67

65

90-95

94

May 26-27

63-70

68

82-86

84

June 3- 4

68-70

69

90-97

94

June 24-25
1922

67-71

70

go-98

94

July 2

68-72

70

It is apparent that air maxima vacillate backward and forward more than
the air minima. In the above tabulation there is a steady climb of the range
average of minima.

This table might permit of the interpretation that Acris might breed at 50°
or 54° or 59° with rains prevalent. But the author believes 62° — 72° or 62°—
69° the optimum or prevalent influential minima in starting vigorous breed-
ing from the last of April to July i.

Egg-laying process. Of the first pair which laid May 15, 192 1, we made
these notes: "Surely the pair doesn't occupy one position but moves about
with each oviposition. It could not have occupied one position above the
stick for eggs to be placed on the stick as they are, i. e., for eggs to drop down
from one position. These were never laid in one mass. Neither are they thus
from the pair breaking up the mass from excessive activity in confined quar-
ters." We did not observe the actual egg-laying but suspect they are somewhat
of the Hyla criicifer type.

On June 4 we have the information that the "two mated pairs of Acris
found June 3, 192 1, laid eggs. One female at least. Believe she laid them
without an embracing male." Three days later June 7 have a similar note
"Female frogs laying without a male. On night of June 3, a female of a
previous pair laid a complement without an attendant male. Previously an-
other female Acris has thus done."

We suspect most of the eggs are normally laid at night. For example, a
mated pair caught the night of June 24 were separated and the female had
laid eggs before morning, and in the field most of the breeding activity comes
at night.

EGGS

Egg moss, egg description. Abbott (1882, p. 707) is the first to speak of
their eggs which "were deposited in httle masses, attached to the blades of
course grass."

Dr. K. V. (Katherine Van Winkle) Palmer who worked over our Acris
collections and the 19 17 transcontinental material found that in an Ames,
(Iowa,) congress of Acris June 12, 1926, that some eggs tended to be in
bunches or masses.

In 191 7 at Dinwiddie, Va., we found this species breeding. Of it we write
in 1920 (p. 29) as follows: "The writer recently found them breeding actively
on June i. They had chosen a shallow (i to 4 inches deep) grassy meadow
pool (PI. X, fig. i). The eggs were attached singly to sedge stems or were

i8o Frogs of the Okefinokee Swamp

strewn singly on the bottom. In one or two instances three or four eggs were
close together. Many of the eggs were in water not more than an inch in
depth. Eggs of Acris hard. Found no more than ten eggs."

In 1921 our first record of eggs came the morning of May 15. "A general
rainy morning. At 6:00 a. m. the mated pair of Acris hadn't laid. At 8:00
there were 74 eggs singly laid. At 8:00 the pair were still mated. At 9:00
when we returned the pair was broken. The eggs all rested on the bottom of
the jar." On the afternoon of the same day "the female of my drinking cup
(male of which leaped out) placed with the three males but she did not lay
single eggs nor was she mated. Nevertheless she laid. Her eggs are in a
single mass on the side of the small bottle in which she was and just above
the water's level." Later the same day we wrote of the first complement
thus: "There are two or three eggs fastened to the stick on its under side. The
eggs are scattered all over the bottom of the jar."

Our field notes on eggs follow: Some Acris gryllus eggs were measured in
the field under adverse circumstances May 16, 1921, "One vitellus was i.o
mm.; outer envelope 3.8 mm. Another had vitellus, i. o mm., inner en-
velope 1.8 mm. and outer envelope 4.0 mm." On May 27, we remark that
eggs laid May 26-27 have "vitellus, i.o mm.; outer envelope 2.8-3.0 mm.; no
inner envelope; egg vitellus creamish white; animal pole deep brown. Must
be something wrong with measurements I made on the eggs of May 16."

On June 25, 192 1 after a pair had laid eggs we examined the eggs in camp
and made notes that the "vitelli were .9 mm., envelope 3.6-3.2 mm., upper
pole 'brownish,' as we have used the term before in egg description (really
nearer buffy olive), lower pole white. Envelope loose, full of trash on outer
edge. Hard to determine where there are two envelopes."

On July 17, 192 1, we for a period mistook singly laid eggs from the bottom
of a pond for Acris eggs. They proved those of Hyla gratiosa which lays
single eggs.

In 1922 on July 3 we artificially secured some eggs from a mated female
which died. Her egg complement was 241 eggs. The eggs taken from the
ovary took up water and the jelly envelope looked as if laid naturally. One
jelly envelope appeared at first but later it very definitely became apparent
that in these ovarian eggs there was a .9 mm. vitellus, an inner envelope
1. 6-1. 8 mm., and a 2.6-2.8 mm. outer envelope."

In Feb. 1923 (p. 34) we merely alluded to Acris gryllus eggs as "single
submerged eggs." Laboratory measurements made by Mrs. Wright and
myself yielded: eggs with vitelli .9-1.0 mm., average .98, mode 1.0 mm. and
envelopes 2.4-3.6 mm., average 2.8 mm., mode 3.0 mm. In one egg an inner
envelope seemed to be present and it was 1.8 mm. in diameter. In 1924
(p. 378) we summarized it thus: "Eggs deposited singly. Envelope single.
No inner envelope or appearance as such, envelope firm, definite in outline
2.4 to 3.6 mm.; vitellus 0.9 to 1.0 mm. Egg complement 241." We are
mindful of two difficulties, one the possibility of egg complement possibly
being laid in a mass occasionally and also of the discovery that an inner
envelope maj^ eventually become the rule rather than the exception.

Acris gryllus i8i

HATCHING PERIOD

The eggs laid May 15, 192 1, between 6:00 and 8:00 a. m. were approach-
ing hatching May 17 at 8:15 when preserved. Air minima of 61°, 64° and 65°
and maxima of 73°, 84° and 76° obtained these three days. The other set of
eggs laid May 15 hatched 12:00 (noon) May 19 or about four days later.
The air mimina from May 15-May 19 ranged from 63°-65° and maxima from
73°-84°, or an average for all of 72°F.

MATURE TADPOLE

Color description from life {July 5, 1921). General coloration dark olive
buff or old gold or ohve lake or sulphine yellow. Belly especially on sides
and on gill region light vinaceous fawn or shell pink or pale salmon color.
Belly with ivory yellow or cartridge buff clusters of spots. These clustered
spots almost cover the top of the body. The region back of the labial mouth-
parts has no light vinaceous fawn nor ivory yellow. The clusters of spots
become almost continuous and look a patch of French gray or lilac gray on
the throat region.

Tail. Tip of tail (upper and lower crests) conspicuously black (at times
lost) . Clusters of ivory yellow or cartridge buff spots on upper crest and upper
part of musculature, less frequent on lower crest and lower musculature.
These spots somewhat amongst black of tail tip. Tail crests almost trans-
parent, certainly translucent almost clear of marks except under lens when
the course of the blood vessels are marked by the body color (dark olive
buff, etc.)

Iris ivory yellow above and below, behind and in front of pupil light coral
red and black. Whole of iris more or less prettily marked with black inter-
spersed with the two lighter colors mentioned.

General appearance. Tadpole medium (42.0 mm.) full and fairly deep
bodied. Tail very long, tail tip very acuminate, with black flagellum. Tail
proportionally the longest and most narrow (depth of tail in length of tail)
of any of the Hylids in the U. S. Dorsal and ventral crests rather narrow,
not equal to the tail musculature in depth. The dorsal crest extends on to
the body to the vertical of the spiracle or just ahead of it but nearer spiracle
than the eye. Spiracle sinistral, mainly directed backwards only slightly
upwards, and it stands out from body in life almost on the lateral axis.
Eye on lateral axis, eye hardly if visible at all from the venter (more like a
Ranid tadpole) suborbital region oblique and vertical as in Hylids in general.
Eye in dorsal aspect just inside the lateral outline. Anus dextral, opening
on a level with the lower edge of the lower crest. Muciferous crypts not
distinct.

Mouth parts: Teeth 2/2. • Upper labium fringed with a continuous row
of labial teeth ; the papillary border does not extend above or inwards beyond
the end of the labial fringe (unlike all our Hylidae of the U. S.). The end of
the second row of upper labial teeth extend beyond the end of the upper
fringe for 1/3 or sometimes about 1/2 of the length of the second upper
labial row (in H. crucifer 1/4-1/6 the length of either lateral row). The

1 82 Frogs of the Okefinokee Swamp

median space between the second lateral upper labial rows of teeth long,
contained 1.2-.75 times in either lateral row, i.e., rarely greater, often equal
to and more often greater than either lateral row. The horny beak is con-
tained in upper fringe i. 2-1. 3 5 times or 1.3 5-1. 5 times in the distance from
one end of lateral row to the end of the other lateral row. There are prac-
tically no inner papillae. The first row of lower labial teeth about equal to
horny beak or shghtly larger. The second row of lower labium usually per-
ceptibly longer than the first row, (the illustration is a little unusual in this
respect) .

Measurements. Length of body (9.6-13 mm.) in tail (26.6-42.2 mm.;
2.5-3.25, average 2.9. Width (5.5-9.0 mm.) of body in its own length 1.44-
1.9, average 1.66. Depth (4.18-8.0 mm.) of body i. 0-1.3 in body depth,
average 1.17. Depth of body i. 625-2.1 in length of body, average 2.0. Depth
(4.2-7.8 mm.) of tail in length of tail 3.25-5.0, average 4.0. Muscular part
(2.8-4.4 mm.) 1. 2-2. 1 5 in depth of tail, average 1.7 1. Spiracle 1.6-2.36
nearer base of hind legs or vent region (3.2-5.6 mm.) than the tip of the snout
(7.0-10.0 mm.), average 1.92. Spiracle to eye usually equal to spiracle to
base of hind legs or vent. Eye i. 0-1.66 nearer the tip of snout (3.0-4.2 mm.)
than to spiracle (4.0-6.0 mm.), average 1.22. Nostril i. 0-1.33 nearer eye
(1.4-2.4 mm.) than snout, (1.8-2.8 mm.). Mouth (1.8-2.4 mm.) usually
1-1.3 larger than internasal space, (1.4-2.6 mm.), average 1.06, rarely less
than internasal space. Mouth contained 1.3-2.2 (average 1.7 1) in inter-
orbital distance (3.0-4.6 mm.). Internasal space contained in interorbital
space 1.33-2.2, average 1.7.

The dimensions of the largest tadpole are:

Total length

42.2

Spiracle to vent

50

Body length

13.0

Spiracle to eye

5-2

Body depth

6.6

Eye to snout

4.2

Body width

8.2

Eye to nostril

2 . 2

Tail length

29.2

Nostril to snout

2.4

Tail depth

70

Mouth

2 .0

Musculature of tail

4.0

Interorbital of distance

4.0

Spiracle to snout

10. 0

Internasal distance

2.6

General remarks. Hay (1892, p. 462) "On the i6th of August . . . found
numerous specimens of the tadpoles of this species. They were found hiding
in the vegetation at the bottom of a small stream. They were in very dif-
ferent stages of development, some with body, the fore and the hind legs
visible, others with only short hind legs." In Indiana, Hubbs (191 7, p. 99) on
Sept. 23 found "tadpoles of species . . . in the pond, as were also a few
transforming individuals."

The first author to describe the mouth parts of the tadpole is 0. P. Hay
(1892, p. 462). He describes them as follows: "The arrangement of the
horny denticles about the mouth of the larvae I found to be different from
that of specimens of Chorophilus triseriatus. In both species there are two

Acris gryllus 183

rows of horny denticles on the upper hp. On the lower lip, there are in
Chorophiliis three rows of denticles, but in Acris only two. Furthermore, in
Chorophilus the denticles are finely serrated at their tips; in Acris this is not
the case. The teeth of the latter genus are less numerous than in the former."
No one else speaks of these mouth parts except Miss Dickerson (1906, p. 156)
who uses Hay's characterization. Had the author of this article remembered
this observation he would not have had so much difficulty in 19 14, 191 7 and
192 1 before he finally identified the tadpoles.

In 192 1 the first time we took adult Acris tadpoles with any degree of
certainty was May 3 1 . From that date to July 5 or 8 we still called them the
black tipped tadpoles with some question as to their identity. For example,
on July 2, we found "tadpoles with black tail tip. Are they Acris or some-
thing else? Not like normal coloration of Acris tadpoles." Even then the
old error of considering H. femoralis tadpoles Acris gryllus still persisted with
us and even to the date of this writing Hyla femoralis tadpoles from 19 1 2-
1 92 1, crop up in our collection labeled .4cm gryllus. It all arose because of a
theory that the alternation of light and brown bands on tail musculature
were transferred to the rear of the femur. Such alternation occurs in the tail
musculature of a Hyla femoralis tadpole and on the rear of the thighs of
Acris gryllus. Hence Hyla femoralis tadpoles were called Acris gryllus
tadpoles for a long time. In fact, even though we were quite satisfied in 1922
we still vacillated per the following note: "July 13, took in an overflow a
quantity of tadpoles. ... A few had a black tip to the tail. Most of
these, however, have lost the black tip. They have a triangle between the
eyes and the banded arrangement on hind legs as do Acris adults. They are
Acris gryllus. But what about the tadpoles recorded earlier as having a light
and dark banded arrangement on the tail, an alternation of bands?" We now
know that the tadpole with alternation of light and dark bands on the tail
musculature is of Hyla femoralis and that the tadpole with a black tail tip
is of Acris gryllus.

This tadpole puzzles us in more ways then one. Of it on July 8, 1921, we
write "Found some tadpoles mature and with black tail tip. These are slim,
long-tailed forms. The most conspicuous thing about them when alive is
the -protruding spiraciilar tube." The tadpole looked like a true frog tadpole
and we were trying to link it up with Rarias whose life histories we did not
know and that went on until the middle of the summer of 1922. Witness the
following: "July 11 in a pond on the island between the first and second dreen
we found in my seine one or two of these long-tailed black-tipped tadpoles.
It was a cypress area, wooded, has Polygala cymosa, Sahbatia decandra,
poison ivy, Hex myrtifolia, Pieris phiUyrei folia, Leucoihae, Clethra. This
tadpole does not have the crest far forward on the back. It has two rows of
teeth above and two below. Can it be Rana virgatipes and that we missed
that species at Billy's Lake in 192 1? It looks more of a Ranid than a Hylid
tadpole. It is not Hyla squirella nor Hyla graliosa.'^ In writing up the
tadpoles of our 1912-1914 we described Hyla femoralis tadpole for Acris
gryllus, but it did not appear in print but unfortunately in 192 1 (p. 33) we

1 84 Frogs of the Okefinokee Swamp

wrote "The conspicuous character is the arrangement of coloration on the
muscular part of the tail. There are four long bands — the first a brownish
band from body to tip of tail; above this a cream white band, followed by
another brown band to tail tip; and this surrounded by another short cream
white band. There are two rows of teeth on the upper lip and three complete
rows on the lower as in the case of some of the frogs." This tadpole de-
scribed is of Hyla femoralis not Acris gryllus. It is one of the mistakes one
assesses to pioneer work.

On Jan. 20, 1923 (p. 406) we corrected this description to ^' Acris gryllus
— very long, black-tipped tails and projecting spiracular tube."

LARVAL PERIOD

Abbott (1882, p. 707) writes 'T did not succeed in following the various stages
of developmental growth from the egg to the matured animal, but was enabled
to determine that it was more protracted than in the case of the common
tree-toad (Hyla versicolor). The difference is, I believe, quite seven weeks."

Hay (1892, p. 462) believed they were breeding March 8 and he ''recorded
transformation Sept. i. This would give 176 days or almost six months,
rather a long period, though twenty-five days or a month in March could be
given to hatching of the eggs."

When we believed Hyla femoralis tadpoles to be Acris gryllus we wrote
(1920, p. 33) "The tadpole of the cricket frog develops in about the same
length of time as that of the peeper. The tadpoles of the former (cricket frog)
transform the same season the eggs are laid and seldom exceed i 1/2 inches in
length." We have had no opportunity of comparing Acris gryllus and Hyla
crucifer tadpoles in the same locality. The above may be true and it may be
false. At that time our evidence seemed to imply 60-80-90 days larval period.

In one pond in 192 1 our first record of breeding comes May 26 and other
breeding notes to June 3. Our first transformation came June 16, 17 and
July 2, July 23. Manifestly the intervals of 13, 23 and 27 days do not seem
reasonable but 59 days from May 2 6- July 23 or 50 days from June 3 -July 23
seem nearer the truth. These June 16 and 17 transformations are from an
earlier breeding so it may have been for the July 2 and July 23 transforma-
tions. In another pond we found great activity on Apr. 24 when a pair was
recorded. The same pond had transformation June 3 and 4 and July 23.
This gives conjectural periods of 41-90 days.

On Billy's Island we have several transformations July 23, 26, 27, also
July 3-12 and June 15. Some breedings in these places May 13 and 12 and
Apr. 24-28 give 32, 33, 48, 52, 59, 60, 66, 70, 70, 71, 75, 79, 86, 90, 90, 94 days
of larval life, or an average of 67 days. Put in another way the first breeding
evidence came Apr. 24, our first transformations were June 3, June 4 and
June 16 or 40, 41 and 53 days but our entry Apr. 22 or 23 into the swamp
precluded earlier records than Apr. 24 by us and these early Acris transforma-
tions were probably from earlier breedings than Apr. 24.

In 1922 we have evidences of 60, 68, 69, 71 and 78 days as possible larval
periods. None of this evidence is positive. Fifty to ninety days seem within
reason as the probable period.

Acris gryllus 185

TRANSFORMATION

Period. Abbott placed transformation in late August. Hay (1892, p. 462)
says "The transformations occurred about Sept i" in Indiana. Hubbs (1917,
p. 99) found a few transforming Sept. 23 in Indiana.

In 191 2 we found transformation likely from May 29- July 15 or later.
In 192 1 from June 3 to July 27, transformation was recorded. In 1922 we
have transformation from June 22-Aug. 29. These transform from Apr. i
or earlier to October i or later.

Size. We recorded in the field little regarding the size of Acris at trans-
formation. On July 7, 1922, we observed that "it has the largest transformed
size relatively (i.e., to adult size) of any of the Anura in the swamp." In 1920
(pp. 34 and 36) we have Acris gryllus transformation size one-half inch and
the adult i 3/4 inches. "The cricket-frog transforms from June i to July if
the eggs be laid early, or in August if the eggs be laid in June. At transforma-
tion it averages a greater length than the swamp cricket frog, being 0.5 inch
in length."

From Gainesville, Ga., as early as Apr. 2, 191 1, J. C. Bradley secured
transformed frogs 10 and 11.5 mm. long, and as late as Aug. 20, 1916, F.
Harper took transformed frogs at Chapel Point, Maryland. These last are
10, 10 and 13 mm. in size. In 1920 we gave one-half inch as the transforma-
tion size. Our 19 12 material yielded transformation sizes of 12 and 13 mm.

In 192 1 on Apr. 28 we found one specimen 13.5 mm. but it was not at
transformation. On June 3 we record one collection with a 13 mm. trans-
formed individual and one 10 mm. with a stub of the tail. Another lot of 5
specimens are 13, 13, 13, 14, 14.5 mm. respectively are marked transformed
but are doubtless slightly beyond. On June 14 we have one transformed at
12 mm. and another 10 mm. with a stub of the tail and a frog mouth. On
June 17 we took one 14 mm. and past transformation. On July 12 we took
three 13, 13.5, 14 mm. beyond transformation. Two transformed at 11.5
and 12 mm. and 3 with sHght stub, these 11.5 and 13 mm. On July 23
we have three just transformed 12, 12, 13.5 mm. On July 2 we found in
one place tadpoles of several sizes and transformations of 10, 12, 12.5 and 14
mm., and one beyond transformation 13.5 mm. On the 24th of July we
found one transformed at 12.5 mm. and one 15 mm. almost transformed,
4 legs, with more frog-like than tadpole mouth. On July 26 we have a
series of six transformed frogs, 11, 11, 11.5 (stub 5 mm.), 12, 12, 13 mm. On
July 27-29 we have one specim.en 13 mm., which is not at transformation.
On the same date the two i i.o and 1 1 .5 mm. just transformed and five others
i3j i3j i3-S> 13-5, 14-5 nim. are possibly just transformed. On July 29 we
have one 11.5 mm. at transformation.

In 1922 on June 27 we took our smallest specimens of transformed frogs,
one 9 mm. in length. On June 29, we have five specimens transformed; lo.o
(tail stub 3 mm.) 11, 11, 12, 12.5 (stub 7.5 mm.) mm. On July 12, at Starling
Branch crossing we secured many mature and smaller tadpoles and trans-
formations 8.5, 10, 10, 10.5, II, II, 12, 12.5 and 14 mm. These are dried up
when measured. On July 14, are seven just beyond transformation 13,

^^Wt>

1 86 Frogs of the Okefinokee Sivamp

i3-5> 13-5) 13-5) 14, 14-5 and 14.5 mm. On July 18 on the prairies we secured
three transformed frogs 12, 13, 13 (tail 6 mm.) mm. On July 28 we have
transformed individuals 11, 11, 11, 11. 5, 12, 12.5 mm. The last has forelegs
just through the skin and the tail 21 mm. long. Finally on Aug. 4, 1922, we
secured two transforming frogs 13.5 (stub 3 mm.) and 14 (stub 15 mm., al-
most transformed frog mouth) mm.

There are 51 specimens in 191 2, 192 1 and 1922 at transformation or
just before. They range from 10-15 mm., 6 at 10 mm. and 2 at 13.5 mm., i at
14 mm., and i at 15 mm. The average is 13 mm. and so is the mode. Thirty-
one of the 51 are 12 mm. or less. We also had 26 debatable stages on individ-
uals possibly beyond transformation. These ranged from 13 mm.-i4.5 mm., 8
at 13 mm., 8 at 13.5 mm., 4 at 14 mm., and 4 at 14.5 mm., or averaging
13.5 mm.

The transformation size usually ranges from 10- 12.5, rarely to 15 mm.
or rarely below 10 to 9 mm. Inasmuch as some tadpoles grow to large sizes
before transformation the larval period must be variable. Furthermore, it is
remarkable to have transformation at 14 or 15 mm. when one male has only
a measurement of 15 mm., and several are clearly males at 17 if not 16 mm.
In other words this species apparently at times almost transforms into
mature breeding frogs.

General remarks. Abbott (1882, pp. 708, 709) in New Jersey records that
"Late in August these tadpoles have become fully developed peepers. Even
then they were very rare during that summer (I suppose this is always the
case) but in September many were found in damp places, never in the water,
but always near a running brook, or a spring. By the middle of September a
marked increase in their numbers was noticed; but their haunts were different.
I found very few in the meadows, but many in damp places, as spring holes,
in the adjacent woodland, and particularly along a brook where the water
flows rapidly over a rocky bed.

"It was here I closed my field studies of these batrachians. Early in Octo-
ber I found a number of these 'peepers' in a little ravine, through which the
above mentioned brook passes. I noticed at this time, that these little
creatures had a decided aversion to the water. Necessary as it was for them
to keep their skins moist, they had no desire to become thoroughly wetted,
and when by chance they made an unlucky jump and settled in the water,
they straightway crawled out and took up a high and dry position on some
projecting stone. If in the sunlight, so much the better. A bath seemed to
chill them, and whenever I drove one into the water, I found that for several
minutes after it emerged I could pick it up without difficulty; but in time
it would regain its ordinary activity, and then quick indeed must be one's
movements who would catch them with the hand alone."

In 191 2 and in the first month of 192 1 our mistaken field identification
of the tadpole made our transformation evidence for these periods too con-
fusing and uncertain to insert here.

In 192 1, June 3, in one pond "Found on lily pads and other surface
vegetation what I at first took for Pseudacris ocularis. The second specimen

Aais gryllus 187

had a stump of a tail and a triangle between the eyes. It is Acris transforming
and transformed. These Acris are about the size of an adult Pseudacris
ocularis. Strangely enough we took a pair the female of which laid when
these tadpoles were transforming. Transformation period must, therefore,
be a long extended one. On June 14 we took transformed frogs at Minne
Lake. On June 16 on Billy's Island in another cypress pond we found all
stages to transformation. On the 17th more transformed individuals were
taken. On July 3 we found many transforming on Honey Island Prairie.
On July 12 more were taken on Billy's Island. On July 23 and July 27
others were recorded.

In 1922 we have several transformation notes: "On June 22 found tad-
poles with large hind legs." On July 2 "on Grand Prairie and whole area
from Lake Seagrove to Grand Prairie are Acris abundant. Found the
species transforming, several on lily pads out from the edge of the islets
('houses, camp houses')." "July 13 at Starling Branch crossing we took in
overflow crossing a quantity of tadpoles approaching transformation. A few
had a black tip to the tail." At the same place "July 24 very few Acris
tadpoles. One with 4 legs, long tail, the stage desired." In another pond
"July 28 found plenty of transformed Acris and a few black tipped tadpoles."
Of another place the same day we wrote "Still a few black tipped tadpoles in
pond near the 2nd dreen."

In August we have a few notes: On Aug. 3-5 on Billy's Island in three dif-
ferent ponds they were transforming. On Aug. 17 at the edge of Starling
Branch crossing "we found a transformed Acris on the ground." On Aug. 31
at two ponds "found several tadpoles with only two legs."

Our field notes above give in 192 1 dates of transformation from June 3-
July 27. In 1922 we have evidences of transformation from June 22 to Aug.
21 and F. Harper later, Aug. 29, found the houses (islands on prairies) "cov-
ered with transformed Acris. '^

In 19 1 2 we secured very little on transformation in Acris gryllus of Oke-
finokee Swamp. Then we rite :

"In some overflow pools on Mixon's Hammock, June 16, 1912, we took
several stages of this form. The pools were about a rod from the west end of
Billy's Lake and had in it also toad and spring frog tadpoles, killifishes and
pigmy sunfishes. Two of the series are at transformation and measure 12 mm.
On May 31- June 2, 191 2, on Honey Island Prairie where water is 4-5 feet
deep we found them common at the surface either fully transformed or with
tails yet visible. These measured 13 mm. These little creatures were resting
on the lily pads of the open prairie or on the vegetation around the little
islands. It is an interesting fact that in these transformed individuals (even
in forms, just before the outpushing of the fore-limbs) the only adult marks
are the conspicuous posterior femoral stripes and rarely the interorbital
triangle. Several other individuals taken at the same time as the trans-
formed individuals measured 16.75-19 mm., and another specimen taken
on Floyd's Island June 25 measured 18 mm. The only mark these eight have
in common is the same femoral coloration. A series of tadpoles taken June 16

1 88 Frogs of the Okefinokee Swamp

represent stages just before outpushing of hind-limbs to transformation and
no doubt the above period outhned for transformation, namely May 2 9- June
16 on this evidence might be pushed to July 15 or later. The breeding must
come early in the spring with that of the spring frog or the development may
be very rapid if the species waits until late April or first of May as some texts
give it. Surely the transformation is not solely in late August or September,
and it is barely possible that the period of final development is shorter than
previously held."

GROWTH

In the Okefinokee region transformation, as we have seen, is from 9-12.5
mm., rarely to 15 mm. The 13 and 13.5 and 14, 14.5 and 15 mm. sizes are
more often beyond transformation than at transformation but 9-15 mm. is its
range. In the north in Maryland and Indiana we have transformations from
10-14 mm., e.g.. Dr. B. W. Evermann at Lake Maxinkuckee, Indiana, took
transformed individuals of sizes 11, 12, 13.5 and 14 mm. In Maryland Mr.
M. K. Brady took it at 12.5 mm., and F. Harper at 10 and 13 mm.

Our Okefinokee males reached 24.5 mm. and the females 24 mm. The
largest males for other parts of its range were 29 mm. and the females 33 mm.
If Hyla squirella transforms at 11-14 mm. and at end of first year as frog
reaches 17-24.5 mm., Acris gryllus a more active frog through the year
ought from 9-15 mm. transformation size to reach in first year old stage 15-
24.5 the largest we have or better to 22 mm. in males and 23 mm. in females.
These first-year-olds would have a mode of 1.8 mm. for the males and 2.1 mm.
for the females. In our study of 169 adults from non-Okefinokee localities we
have a range considerably beyond 24.5 mm. our largest in the Okefinokee.
Some of these reach 33 mm. (U.S.N.M. 3935). The lowest and highest modes
for these males are 20 mm. and 25 mm. and for females 22, 24 and 28 mm.
Possibly in the northern localities where transformation sizes are the same as
for our smaller Okefinokee adults and where the growing season is not so
long frogs from 23-29 mm. males or 26-33 i^i^- females may be two-year-olds.

We, therefore, postulate somewhat reasonably the following: transforma-
tion 9-12.5 mm. (rarely to 15 mm.); ist year olds, 15-22 or 23 mm.; possible
two-year olds, 22 or 23-29 to 7,7, mm.

AUTUMNAL DISAPPEARANCE

Abbott (1882, p. 710) in 1881 in New Jersey writes: "Sensitive as these
'peepers' are to changes of temperature, it is by no means the first frost that
drives them into their winter quarters. In the autumn of the past year (1881)
I found them last as late as Nov. 12th but even later (Dec. 27th) my son
found one in the meadows which was as liyely as a cricket. The frogs gen-
erally were singing this day. For more than two weeke prior to Nov. 12th
there had been several white frosts, and the true frogs (Ranae) had all dis-
appeared except su&h few as lingered in the warm waters of the larger springs."

In Illinois H. Garman (1892, p. 342) finds that "Examples nearly grown
were taken November 17, 1888, under logs in the vicinity of a creek in Cham-
paign County, where they were hibernating."

Acris gryllus 189

In June, 1896, C. S. Brimley (p. 501) makes the "cricket frog abundant,
active all the year round except in the severest weather." Ruthven, Thomp-
son & Thompson (1912, p. 46) finds "This species hibernates during the cold
weather but soon becomes active again during warm periods." In Colorado
M. M. Ellis and J. Henderon (1913) find it until Oct. 26. In Virginia, Nelson
County, Dunn (1916, p. 22) finds one Nov. 26. Brimley has taken it at Bay
St. Louis on Dec. 4 and at Raleigh on Dec. 12.

In 1913 and 1914 the Cornell party of Dec. 22, 1913-Jan. i, 1914, found
this species active and took several specimens. Doubtless this species is
more or less active throughout the year in the Okefinokee Swamp.

FOOD

Holbrook (1842, Vol. IV, p. 132) gives this note: "It feeds on various
kinds of insects, and makes immense leaps to secure its prey, or to escape its
pursuers. It can easily be domesticated, and takes its food readily from the
hand; I have kept several for months in a glass globe on a few sprigs of
purslain {Portulacca olemcea), feeding them occasionally with flies."

Abbott (1882, p. 708) finds that "They fed ravenously at this time (breed-
ing season) and even when confined in very cramped quarters, would devour
any flies that came within reach." Later (p. 710) he states that "My im-
pression is, that they do not require or partake of any food during their brief
existence as matured 'peepers' in autumn (i.e., from completion of growth of
their limbs in September to the commencement of their hibernation). My
reason for this is based upon the fact that the specimens in a bottle, to which
I referred, were placed in confinement on the 20th of October, 1881, and the
date of writing, Jan. 29, 1882, a period of one hundred days has just elapsed.
During this time these 'peepers' have had no food, have been quite as active
as their limited quarters would permit, and yet have not lost weight to any
important extent. One which I weighed on the day following its capture
weighed forty-four grains and seventy-five days later had lost but one grain
in weight. It is very different in spring; then they are voracious feeders and
capture millions of minute insects. At this time their stomachs are always
full; and while the size of the animal is not larger than in autumn, the weight
is nearly twice as great."

To believe that they feed not from transformation to autumnal disap-
pearance is foreign to the facts.

H. Garman (1892, p. 342) gives some definite items. "Its food consists of
insects, and if the habits of the frogs led them more frequently into cultivated
grounds they would doubtless do good service to agriculture in destroying
aphides. Among other insects, Chlorops, crane flies, Thyeocoris, Calocoris
rapidus, numerous pupae and wingless female Aphididae and Orthoptera
have been determined from the contents of their stomachs."

ENEMIES

On 19 1 2 specimens and collections we made the following observations:
"It appears from stomach examinations that the enemies of Acris gryllus

I go Frogs of the Okefinokee Swamp

are as much aquatic as terrestrial. The snake which had eaten the cricket-
frog was the aquatic southern riband snake (Tham?iophis s. sackeni) and
none of the numerous specimens of black or hog-nosed snakes has eaten it,
strong negative evidence indicate that it is quite aquatic in nature. No
doubt it proves one of the best sources of food for the herons, ibises and
cranes."

In 192 1 we have the following evidence. During a congress of .4cm
Apr. 25 in one pond Thamnophis sackeni was on the surface and ever present.
On July 27 "on the prairie just north of Black Jack Island (F. Harper) came
upon a seven-inch ribbon snake {T. sackeni), which had a grip on the hind
leg of an Acris, . . ."

Other notes are on Apr. 28 we found fishermen who used Acris for "perch"
bait. "Some think them better than crawfish." "Others used them for set-
line bait." On May 6 we found "a cypress pond where I caught so many
Acris earlier. They must have been leaving the pond before. This time there
were few of them. The pond was almost dried up. Around little water re-
maining, in the center was a mud flat covered with dried and drying
aquatic vegetation. Under this mat were young Rana grylio, Acris, etc.
A pig was rooting up these frogs."

"Florida Crackles and a Great Blue Heron were walking around in the
little center pool after Elassoma, water insects, frogs and tadpoles."

Abbott (1882, p. 708) in contrast to cricket frogs' ravenous appetite
writes "On the other hand, they were the main food supply of certain fishes,
all the snakes, the turtles and a few species of birds. I find that all of our
snakes at this time (April and May) were more abundant in the meadows
than elsewhere, and have no doubt (been) drawn thither for the purpose of
feeding on these little batrachians. Even that lover of high, dry and dusty
fields, the hog-nosed snake ( Heterodon platyrhinus) was found to be stationed
at intervals along the ditch banks, on the lookout for 'peepers'; the dis-
section of one of these snakes proved that it had fed upon these small frogs."

In 191 7 Hubbs (p. 99) describes in considerable detail an interesting one-
legged cricket frog "whether this monstrosity is the result of a mutation or of
a very early injury can not be stated."

AFFINITIES

LeConte, its describer, called it Raria gryllus. Harlan (1835, pp. 104, 105)
repeats his description, holds it erroneously supposed by Daudin to be the
young of the Hyla lateralis, and gives its length as "one and one-half inches."

Harlan further describes (p. 105) another form Rana dorsalis. "Char. —
Above fuscous, smooth, with a broad, green or reddish, longitudinal vertebral
band, bifurcating anteriorly, and extending over each eye; snout above, pale
or whitish; beneath white, throat and inner part of the thighs freckled; a white
line on the side of the neck, extending from the eye to the scapula. Length of
the body 8/10 of an inch; of the legs i 1/2 inches. This measurement being
taken from the largest of seven specimens. In habits Florida, Carolina and
New Jersey. May prove a variety of R. gryllus. . . ."

Acris gryllus 191

Figured at p. 72 of this volume. This is very apparently Acris gryllus
and was really first described by Harlan in 1826 (p. 340).

In 1841 Dumeril and Bibron (1841, pp. 507-509), place it and Pseudacris
nigrita right after the genus Litoria as the two species of Acris. Specimens of
each of these "deux tres-petites especes" they received from LeConte.

Holbrook's (1842, Vol. IV, p. 133) "General Remarks" about Hylodes
gryllus are "The first notice of this animal is found in Bartram's Travels in
CaroHna and Florida; he calls it the Savannah Cricket, and gives a tolerable
account of its manners. This animal forms a good connecting link between
the genera Rana and Hyla, and seems to partake of the habits of both, for
though it hkes water at all times, yet it may be found on aquatic plants
waiting for its prey ; and it has the power of adhering to smooth surfaces like
the Hylae, but much less perfectly, as it can never sustain itself on the under
surfaces of leaves, etc."

"For the present I am disposed to arrange this animal with the Hylodes
of Fitzinger: for i. The extremities are slender, with the fingers and toes only
slightly swelled at the tips; and 2. It cannot belong to the genus Rana, as it
wants a bony sternum, etc."

"Dumeril and Bibron have established a new geuns Acris, for the reception
of this animal and the Cystignathus nigritus; but I see no reason for removing
it from that of Hylodes, especially as their genus is characterized by having
the toes free." In 1849 (Appendix, p. 15) Holbrook still retains it in Hylodes.
DeKay (1842, Part III, p. 70) "followed Dr. Holbrook in arranging it under
Hylodes."

In 1855 to 1858 we have more activity in Acris species than at any other
period unless it be the gryllus-dorsalis period of 1825-26. In 1855 Baird
(PP- 59> 60) gives two forms as follows "Acris crepitans, Baird. — Brownish
above. The median region of head and body above bright green; a dark
triangle between the eyes. Three oblique blotches on the sides, nearly equi-
distant, the first behind the eye, the last on the flanks and running up the
back; all usually margined with lighter. A narrow white line from the eye to
the arm. Beneath yellowish white. Inferior face of thigh plain. Tibia a
little more than half the length of the body; foot rather smaller. Head
rather obtuse, scarcely longer than broad. Web of hind foot extending to the
penultimate articulation of the 4th toe. Syn. Hylodes gryllus DeKay, N. Y.
Zool. Ill, (1842) 70, PI. XXII, f. 61. Hab. Northern States generally.
"2. Acris acheta Baird. — Slender. Tibia two-thirds the length of body.
Color above dark brown; blotched much as in Acris gryllus. Beneath white,
closely mottled with brown on the body and on the inferior and inner faces of
the limbs; where are also visible minute dots. Lower surface of body appears
quite black. A narrow white line from eye to arm. Hah. Key West, Florida."

The first writer who attempts minutely to distinguish Acris gryllus and
Acris crepitans is LeConte (Dec. 1855, pp. 426, 427) the first describer of the
former. We will not copy his descriptions but he says of the latter (A. crepi-
tans) that it "very much resembles the preceding (A. gryllus) but the head is
shorter and blunter, the body likewise is not as slender or as graceful in form

1 92 Frogs of the Okefinokee Sivamp

as in A. gryllus having a considerable resemblance to a toad. Varies ex-
tremely in color and in other marks. The only unvarying marks which I have
been able to disover are the warts on the back, the dusky line from the axilla
on the side (which sometimes almost vanishes), the darker angular spot above
the anus (which at times is scarcely perceptible) and the white granules on
the hinder and under parts of the thighs. . . ."

In 1858 (pp. 70, 71) Gunther places the genus "Acris Disk small. Toes
broadly webbed; fingers free; tongue heart-shaped. North America." in the
Polypedatidae. Interestingly enough he places Acris picker ingri in this
genus and holds "This genus might belong to the family of Hylidae. Bibron
and also Baird describe the diapophyses of the sacral diapophyses of the
sacral vertebra as not dilated ; but I ask naturalists having the opportunity
of examining fresh specimens to direct their attention to this subject. The
relation to Pseudacris nigrita is very great, which species also exhibits only
slightly dilated processes."

In 1882 (pp. 336, 337) G. L. Boulenger gives three forms. "Acris gryllus. —
United States: the typical form confined to the Austral riparian region, the
var crepitans to the Eastern and Central regions."

"Var crepitans — Differs by a stouter habit, the head being shorter, and the
tibio-tarsal articulation reaching hardly the tip of the snout. According to
Mr. Cope (Check-list N. Amer. Rept.) this form should not be specifically
separated from the preceding, but both are considered subspecies."

Var. hufonia. Resembles the preceding, but the head and back are covered
with very large warts. Markings indistinct. Perhaps a species. 9 New
Orleans.

Cope (1889, pp. 324, 325) holds that "It is quite possible that it may
become necessary at some future time to unite this genus with Hyla." . . .
The northern and southern sections of this area produce forms which offer
considerable differences, but which must be termed subspecies, on account of
the existence in some localities of intermediate individuals. I have seen
such from lUinois, Pennsylvania, Missouri, and elsewhere.

"These subspecies are as follows: Hinder foot less tarsus less than half of
the length of the head and body; dermal tubercles larger; posterior femoral
stripe less distinct — Acris g. crepitans. Hinder foot less tarsus longer than
half head and body; dermal tubercles smaller; femoral stripe very distinct. —
Acris g. gryllus.

"These subspecies are respectively of northern and southern distribution,
the A. g. gryllus ranging from North Carolina to Florida and Louisiana."

"Specimens from the lower Mississippi are frequently of obscure colors,
of rather larger size, and with large tubercles. I have not been able to dis-
tinguish them as forming a constant subspeeies. Mr. Boulenger . . . refers
such a specimen to a *var. bufonia.' "

"As regards the transition of this subspecies and the A. g. crepitans, a
number of specimens display intermediate proportions. Thus in some the
posterior foot, minus the tarsus, is exactly half the length of the head and

Acris gryllus 193

body. In some lots from a single locality some specimens have the hind foot
one-half the length, while others fall a little below and still others fall a little
above this proportion.

"The distinctness of the posterior femoral brown stripe is subject to vari-
ation. . . ."

In 1892 Hay considers Acris "Closely allied to Hyla," and gives the two
subspecies as occurring in Indiana just as E. E. Ramsay (1901, p. 222), and
McAtee (1907, p. 15) do for Monroe County, Indiana. H. Garman (1892, pp.
342, 343) holds "LeConte's characterization of the two forms . . . the
best extant, but the only difference he presents which in so variable a species
is of a varietal importance, is the size (1.4 inches for variety gryllus and 1.2
for variety crepitans). None of the Illinois specimens examined are more
than 1.25 inches in length of body."

S. N. Rhoads (1896, pp. 396, 397) adds more to the gryUus-crepitans dis-
cussion "Examination of nearly forty specimens from widely separated
localities in Tennessee and Kentucky fails to reveal any constant distinctions
between gryllus and crepitans, if both forms are found in the State. Prof.
H. Garman considers the size of gryllus (1.4 in.) as given by LeConte as the
only reliable difference. This is much larger than any in my series, and as
the Samburg specimens are smaller and more slender than those from the
Cumberland plateau, the theory that the Southwestern frogs are larger
than Northeastern ones is contradicted. LeConte defines the habitat of
gryllus in the South Atlantic States, and that of crepitans in the remainder
of the eastern United States, but Prof. Cope's identication of the Smith-
sonian series allots specimens of both forms to both sides of the Allegheny
Mountains in such a way that faunal definitions lose their significance. Such
being the case I have lumped the entire Tennessee series under the original
specific name." W. P. Hay (1902, p. 128) holds gryllus and crepitans "of
very doubtful vahdity." Deckert (1915, p. 22) observes that "This species
looks and acts more like a water frog than a tree-toad, although belonging to
the latter family." In 191 7 Dunn (p. 621) finds his material from the North-
west of North Carolina intermediate between A. g. gryllus and A. g. crepitans,
but in 1923 Brimley and Mabee (1925, p. 15) find in eastern North Carolina
both Acris crepitans and Acris gryllus.

Earlier than 1923 Percy Viosca, Jr., believed he could separate Acris
gryllus and Acris crepitans. In 1923 (p. 43) he discusses this matter thus:
"The puzzling status of the Genus Acris, as far as Louisiana is concerned, has
been positively cleared by these studies. There are two distinct species in
Louisiana, the upland species being, tentatively, Acris gryllus, and that of the
lowlands, Acris crepitans. Wherever their ranges overlap, they are found side
by side without interbreeding, each with its characteristic chorus and habits."

In 1925 we took one evening trip with him in February but the
author did not get enough experience in the one evening to warrant his at-
tempting to tell them by song, habits and structural and color characters.
If any one can do it Mr. Viosca is the person with experience to do it. Fur-
thermore, Mississippi Valley, Texan and southeastern forms crowd into

194 Frogs of the Okefinokee Swamp

Louisiana. From New Orleans Boulenger described his "var. hufonia."
This region ought to have a diversity of Acris forms.

It is interesting that this form was described as a Rana. In the general
appearance of its tadpole we considered it quite Ranid-like. In fact, for a
time until we came to know Rana virgatipes tadpoles we wondered if these
Acris tadpoles might be the young tadpole of Cope's frog. Gunther put
Acris gryllus and Acris pickeringii {Hyla crucifer) in the same genus. Each
lays single eggs with a single envelope though in each there are some evi-
dences of an inner envelope and each has vitellus .9-1.0 or i.i mm.

Dumeril and Bibron established Acris for Acris gryllus and Acris nigritus
(Pseudacris nigrita). Normally the various forms of P. nigrita lays eggs
in masses not singly but Abbott generally and we rarely recorded masses for
Acris gryllus. Pseudacris ocularis lay very similar single submerged eggs with
a single envelope 1.2-2.0 mm., as in Acris gryllus. The Pseudacris iriseriata
or P. nigritus triseriatus tadpoles of western New York normally have teeth
formulae of 2/2 like Acris gryllus. Holbrook placed Acris gryllus and Pseudacris
ocularis, each in Hylodes.

In eggs it might be linked with Pseudacris ocularis, Hyla crucifer,
Hyla squirella or Hyla gratiosa. In tadpoles it is in some ways apart from
all the Hylids of the U. S. A. In other ways it falls close to Hyla crucifer
or some species of Pseudacris. In transformation size it falls in with Hyla
squirella, Hyla andersoni and Hyla femoralis. In tadpole coloration it is
apart. Acris is sufficiently distinct to be retained yet awhile although it
has characters in common with Pseudacris and Hyla. It is the most dis-
tinctive and commonest Hylid in U. S. A.

We have made no endeavor to separate Acris gryllus and Acris crepitans.
They may be distinct. The only person who has given much attention to this
question in recent times is Mr. Percy Viosca, Jr., who believes them distinct
in appearance, habits, call, etc. Possibly he is right.

BIBLIOGRAPHY

1882 Abbott, C. C. Notes on the Habits of the Savannah Cricket Frog. Am. Naturalist

XVI, 1882, pp. 707-711.
1896 Atkinson, C. Batrachia of Turkey Lake, Ind. Proc. Ind. Acad. Sci., 1895, Feb.

1896, p. 258.
1892 Blatchley, W. S. Notes on the Batrachians and Reptiles of Vigo County, Indiana.

Journal Indiana Soc. Nat. Hist., XIV, 1891-1892, p. 27.
1882 Boulenger, G. A. Catalogue of the Batrachia Salientia — . British Museum, 2nd

edit., London, 1882, pp. 336, 337.
1914 Boyle, H. S. Some Notes on the Cricket Frog on Long Island. Copeia, June 20,

1914 no. 7, p. 4.
1896 Brimley, C. S. Batrachia Found at Raleigh, N. C. Am. Nat. 30, 1896, p. 501.
1925 Brimley, C. S. and Mabee, W. B. Reptiles, Amphibians and Fishes Collected in

Eastern North Carolina in the Autumn of 1923. Copeia, Feb. 16, 1925, No. 139, p.

15-
1889 Cope, E. D. The Batrachia of North America. Bull. U. S. Nat. Mus., No. 34, 1889,

pp. 324-329-
1892 . The Batrachia and Reptilia of Northwestern Texas. Proc. Phila. Acad. Sci. ,

Vol. 44, 1892, pp. 333.

1914 Deckert, R. F. List of Salientia from near Jacksonville, Florida. Copeia, Feb. 14,
1914, No. 3, p. 3-

191 5 . Concluding Notes on the Salientia of Jacksonville, Florida. Copeia, July

27, 1915, No. 20, pp. 21-22.

Acris gryllus 195

1849 DeKay, James E. Zool. of N. Y., Part III, p. 70.

1902 Dernehl, P. H. Place-Modes of Acris gryllus for Madison, Wise. Bull. Wise. Nat.
Hist. Soc., Vol. 2, n.s., No. i, January 1902, pp. 75-82.

1906 Dickerson, M. C. The Frog Book. pp. 15,^-156.

1905 Ditmars, R. L. The Batrachians of the Vieinity of New York City. Guide Leaflet
No. 20, Am. Mus. Journal Vol. 5, pp. 192, 193.

1841 Dumeril, A. M. C. and G. Bibron. Erpetology — Paris, Vol. 8, pp. 507-509.

1916 Dunn, E. R. Notes on Virginia Herpetology. Copeia, Mareh 24, 1916, No. 28, p. 22.

1917 . Reptile and Amphibian Collections — North Carolina Mountains — . Am.

Mus. Nat. Hist. Bull. No. 3, Art. 23, p. 621.

1913 EUis, M. M. and Junius Henderson. The Amphibia and Reptilia of Colorado.
Part I, Univ. Colo. Studies, Vol. X, No. 2, May 13, pp. 58, 59.

19 1 5 . Part II, Vol. XV, No. 6, June 19 15, p. 258.

191 7 . Amphibians and Reptiles from the Pecos Valley. Copeia, Apr. 24, 19 17,

No. 43, p. 39.

1907 Fowler, H. The Amphibians and Reptiles of New Jersey. Annual Rept. N. Y.
State Museum, 1906, 1907, pp. 99-104.

19 14 Gaige, H. T. A List of Amphibians and Reptiles Observed in Richland County,
Illinois, in May 1913. Copeia, Oct. 15, 1914, No. 11, p. 4.

1892 Garman, H. A Synopsis of the Reptiles and Amphibians of Illinois. Bull. 111. State

Lab. Nat. Hist., Vol. Ill, Art. XIII, 1892, pp. 340-343.
1858 Gunther, A. Catalogue of the Batrachia Salientia — British Museum, London,

1858, pp. 70, 71.
1826 Harlan, R. Journ. Acad. Na,. Sei., Vol. V, p. 340.

1835 . Medical and Physical Researches, Phila. pp. 104, 105.

1892 Hay, O. P. The Batrachians and Reptiles of the State of Indiana. Ind. Dept.

Geol. Nat. Resources, 17th Annual Rept., 1891, Indianapolis, 1892, pp. 461-463.

1892 Hay, W. P. A List of the Batrachia and Reptilia of the District of Columbia and
Vicinity. Proc. Biol. Soc. Wash., Vol. XV, June 20, 1902, p. 128.

1842 Holbrook, J. E. North American Herpetology, Vol. IV, 1842, pp. 131-133.
1849 . In George White's Statistics of Georgia 1849 Appendix.

1917 Hubbs, C. L. A One-Legged Cricket Frog. Copeia, Oct. 26, 1917, No. 50, p. 99.

1893 Hurter, Julius. Catalogue of Reptiles and Batrachians found in the Vicinity of
St. Louis, Mo. Trans. Acad. Sci. St. Louis, Vol. VI, No. 11, Dec. 12, 1893, p. 253.

1825 Le Conte, John. Remarks on the American species of the Genera Hyla and Rana.

Lyceum Nat. Hist. N. Y., Vol. I, May 16, 1825, pp. 278-282.
1855 . Desc. Cat. of the Ranina of the United States, Proe. Acad. Nat. Sci. 7,

(1854, 1855), 1856, pp. 426, 427
1895 Loennberg, E. Notes on Reptiles and Batrachians Collected in Florida in 1892

and 1893. Proc. U. S. N. M., Vol. XVII, 1894, 1895, P- 3.38.
1925 Logier, E. B. Shelley. Notes on the Herpetology of Point Pelee, Ontario. The

Canadian Field Naturalist, Vol. XXIX, No. 5, May, 1925, p. 92.
1907 McAtee, W. L. A List of Mammals, Reptiles and Batrachians of Monroe County,

Indiana. Proc. Biol. Soc. Wash., Feb. 25, 1907, Vol. XX, p. 15.
1 891 Morgan, T. H. Some Notes on the Breeding Habits and Embryology of Frogs.

The American Naturalist, Aug. 1891, Vol. 25, pp. 753-760.
1904 Morse, Max. Batrachians and Reptiles of Ohio. Proc. Ohio State Acad. Sci., Vol.

IV, Part 3, Special Paper No. 9, June 5, 1904, p. 118.

19 14 Overton, F. The Frogs and Toads of Long Island. Mus. Brookl. Inst. Arts and
Sciences, Science Bull. Vol. 2, No. 3, Nov. 3, 1914, pp. 30, 31.

1901 Ramsay, Earl E. The Cold Blooded Vertebrates of Winona Lake and Vicinity.

Proc. Ind. Acad. Science, 1900, 1901, p. 222.
1895 Rhoads, S. N. Contributions to the Zoology of Tennessee, No. i, Reptiles and

Amphibians. Proc. Acad. Nat. Sci. Phila., 1895, 1896, pp. 396, 397.
1912 Ruthven, A. G. The Herpetology of Michigan. Mich. Geol. and Biol. Survey,

Publ. 10, Biol. Ser. 3, 1912, pp. 45-47.
1898 Sherwood, W. L. The Frogs and Toads Found in the Vicinity of New York City.

Proc. Linn. Soc. N. Y., 1897-1898, No. 10, pp. 18, 19.

1903 Stone, W. Proc. Acad. Nat. Sci. Phila., Vol. 55, Aug. 1903, pp. 538-539.

1910 Strecker, J. K., Jr. Notes on the Robber Frog (Lithodytes latrans Cope). Trans.
Acad. Sci. St. Louis, 1910, Vol. 19, pp. 73-82.

19 1 5 . Reptiles and Amphibians of Texas. Baylor Bulletin Vol. XVIII, No. 4,

Aug. 1915, p. 49.

1923 Viosca, Percy, Jr. An Ecological Study of the Cold Blooded Vertebrates of South-
eastern Louisiana. Copeia, Feb. i, 1923, No. 115, p. 36, 43.

1920 Wright, A. H. Frogs: Their Natural History and Utilization. Rept. U. S. Com.
Fisheries 1919, Appendix IV, Doc. 888, 1920.

1923a . The Tadpoles of the Frogs of Okefinokee Swamp, Georgia. The Anat.

Record Vol. 24, No. 6, Jan. 20, 1923, p. 406.

ig6 Frogs of the Okefinokee Swamp

1923b . The Salientia of the Okefinokee Swamp, Georgia. Copeia, Feb. i, 1923,

No. 115, p. 34-
1926 Wright, A. H. The Vertebrate Life of Okefinokee Swamp in Relation to the Atlantic

Coastal Plain. Ecology, Jan. 1926, Vol. VII, No. i, p. S:i.
1924 Wright, A. H. and A. A. Wright. A Key to the Eggs of the Salientia East of the

Mississippi River. The American Naturahst, Vol. LVIII, July-August, 1924, p. 378.

Pseudacris nigrita (LeConte)

(PI I. Fig. 9)
COMMON NAMES

Swamp Cricket Frog. Swamp Tree Frog. Swamp Chorus Frog. Swamp
Tree-toad. Striped Tree-frog. Rough Chorus Frog. Black spotted Tree Frog.

RANGE

Check list. "Type locahty not given. Range: South CaroHna to Mis-
sissippi." Stejneger & Barbour 1923, p. 28.

Supplementary records. C. S. Brimley (19 10) has "Chorophilus nigritus
Rough Chorus Frog" from Bay St. Louis, Mississippi, Feb. 10, 1898, i."
At Jacksonville, Fla., R. F. Deckert (1914, No. 3) found "Chorophilus
nigritus LeConte, common during the winter months." Fowler (191 7, p. 39)
records it at Brickells Hammock and Mersa Isle near Miami, Fla., in Feb-
ruary and March 19 16. Loding (1922, p. 17) records it in Mobile County,
Ala. In South Carolina at Greenville A. L. Pickens (1927, p. 109) secured
several specimens. Holbrook (1842 Vol. IV, p. 108) gave its distribution
as Georgia and Carolina. LeConte observed it in the former State, and I
have seen it in the latter. LeConte (1855, p. 427) holds it "Inhabits Georgia
and South Carolina . . . " Our records and others like Cope send it into
Florida. 0. C. Van Hyning at Gainesville, Florida has sent me several times
representatives of this species.

Local Okefinokee records. Our records are wholy outside the swamp
along its eastern border and at Fargo, Ga. On June 15, 1922, M. D. Pirnie
and the author took two specimens two miles east of Chesser Island. The
next was taken south of Spanish Creek near Folkston-Moniac road July 15,
1922. Three days later we took one at Murray Bay northeast of Chesser
School and Starhng Branch. On August 16, 1922 we secured one at one to one
and a fifth miles south of HilUard, Florida. In 192 1, August 5, F. Harper
recorded it at Fargo, Ga., on the southwestern edge of the swamp. In 1909
September 11, Professor J. C. Bradley collected a Pseudaci is which at the
time we identified as Chorophilus occidentalis. It looked to be P. nigrita and
comes from near the Okefinokee Swamp. This same collector secured two
P. nigrita at Tifton, Ga., September 8, 1910.

GENERAL APPEARANCE

On May 16, 1825 Captain John LeConte characterizes this form as fol-
lows "6 Rana nigrita; above black, speckled with small white warts; middle of
the back cinereous with an interrupted stripe of black, upper lips with a white

Pseudacris nigrita 197

line; beneath granulate whitish; sides golden; legs barred with whitish, hind
part of the thighs brown; hind legs very long." If, for example, a specimen
be faded out one of the outstanding characters will be the last one of long legs
in LeConte's preliminary "indication" or description.

Thirty years later (1855, p. 427) this author describes it at length —
"Chorophilus nigrita .... Body elongated rather narrow, above with
numerous small warts. . . . Tympanum distinct, black or of the color of the
body. . . . Beneath . . . (in dead specimens appearing granulate) . . . ;
thighs and tibiae granulated, more or less covered with small warts. A
very slightly perceptible web between the second and third and between the
third and fourth toes, the fourth and fifth are closely united at the base.
Fingers and toes with small dilations at their tips. . . ."

In 1 84 1 (p. 509) Dumeril at Bibron gives it "Caracteres. Deux groupes de
dents vomeriennes affectant une forme en chevron. Tympan assez distinct.
Une tres-courte membrane a la racine des orteils."

In 1862 Holbrook (Vol. IV, p. 107) gives "The head is elongated, . . .
the snout rather pointed. The nostrils are lateral, and nearer the snout than
the orbits. The eyes are large and prominent .... The mouth is rather
large, and the palate is armed with two thick groups of minute teeth, placed
between the posterior nares and disposed a little 'en chevron'. . . . The
body is slender .... The anterior extremities are slender . . . ; there are
four long and delicate fingers. The posterior extremities are very long . . . ;
there are five toes, not palmated."

DeKay (1842, Part III, p. 65) in his Extra-limital forms of New York
gives "Cystignathus nigritus (Id. Holbrook Vol. 3, pi. 19; and Vol. 4, pi. 26,
2nd Ed.) small, olive brown; an interrupted black vertebral line, with black-
ish blotches along the sides; legs barred; upper lip white. Length i3^ inches.
Carolina, Georgia."

In 1858 A. Gunther (p. 97) gives its color as "Blackish ash, with three or
five darker, sometimes interrupted longitudinal bands; a black streak passing
through the eye above the tympanum; upper lip with a whitish streak, a.
Adult. Georgia. From Mr. Cumming's Collection, b. Adult. Great Bear
Lake. Presented by Sir J. Richardson." Of course, this last refers to P.
septentrionalis.

A. L. Pickens (1927, p. 109) records that "Specimens from A. L. Pickens,
Greenville (S.C.) are indefinably intergraded with some other forms. To
me some appear to have inclination toward P. triseriata, others, perhaps
toward P. feriarum. The more typical form seems to appear in the lower
Piedmont."

COLORATION OF SPIRIT SPECIMENS

LeConte's first extended description of its color comes in 1885 (p. 427): —
Color above dark slate with more or less of oblong or oval or round, some-
times very numerous black spots, which often form stripes. Upper lip white-
which color often extends to the insertion to the arm. I rids golden .... A
broad black band extends from the nose through the eyes and along the sides

198 Frogs of the Okefinokee Swamp

to beyond the middle of the body. Beneath whitish, sometimes tinged with
yellow with a few black spots irregularly assembled in groups, cancellately
reticualate . . ., arms and legs spotted and barred with black .... Chin
often dusky.

(We will not undertake to describe coloration in spirits. See coloration
from life under mating).

STRUCTURAL CHARACTERS
(See General Appearance)

Cope (1889) held that the skin was warty, above the head acuminate, the
width entering the total three times; heel reaching anterior to orbit; size
larger. All these characters are contrasted with C. feriarum and C. triseriatus.
Hay in 1892 follows the same general characterization of C. nigritus nigritus.

Baird (1854, p. 60) when he separates Chorophilus nigritus from Helo-
caetes triseriatus, H. clarkii and H. feriarum he said it was "smooth above,
granulated beneath. Extremities of limbs simple not dilated at tip. Hands
free; feet with a slight basal web, which is entirely wanting between the two
other toes. Tongue orbicular; emarginate behind. Teeth posterior to the
nares. Tympanum distinct. Transverse apophyses of sacral vertebrae
dilated into triangular pallets."

MEASUREMENTS

(Recent Material)

Head to angle of mouth 1.15 (25 mm.cf)-i.o, (28 mm.9) in width of
head; head to rear of tympanum 1.0-.9 in width of head; head to angle of
mouth 3.37-3.1 in length of body; head to rear of tympanum 2.95-2.8 in
length of body; snout .52-.87 in first finger; snout .74-.87 in fourth finger;
snout .60-.76 in first toe; eye i. 50-1. 2 in snout; eye .60-54 in tympanum;
eye .8-.87 in first finger; tympanum 3.4-3.4 in intertympanic width; tym-
panum 2.5-1.84 in snout; internasal width .80-.83 in upper eyehd width;
interorbital width .80-.83 in upper eyelid width; interorbital width i. 0-1.2 in
internasal width; interorbital width 2.8-3.4 in intertympanic width.

Forelwib. Fore hmb — 1.92-2.0 in length of body; fore limb 3.0-3.0 in
hind limb; first finger 2.0-1.75 in third finger; second finger 1.7-1.4 in third
finger; second finger .85-.8 in first finger; third finger .70-. 71 in second toe;
fourth finger i. 4-1. 7 5 in third finger; fourth finger .70-1.0 in first finger;
internasal width 1.1-1.33 in first finger; internasal width i. 27-1. 66 in second
finger; internasal width 2.18-2.33 in third finger; internasal width 1.5- 1.3 3
in fourth finger.

Hindlimh. Length 1.5 6- 1.5 in hind limb; tibia 2.0-2.0 in length; tibia
3.1-3.0 in hind limb; tibia i. 03-1.0 in fore limb; tibia .9-. 95 in hind foot
without tarsus; first toe 1.2 1- 1.4 in second toe; first toe 2.57-2.4 in third toe;
first toe 2.71-3.6 in fourth toe; first toe 2.64-2.6 in fifth toe; second toe 2.1 -1.9
in third toe; second toe 2.2-2.6 in fourth toe; second toe 2.1 -1.8 in fifth toe;
third toe 1.05- 1.3 5 in fourth toe; third toe 1.02 -.9 5 in fifth toe; fourth toe
1. 5-1.0 in hind foot; fourth toe i. 65-1. 06 in tibia; fourth toe 1.71-1.06 in fore

Pseudacris nigrita 199

limb; fifth toe i. 02-1. 44 in fourth toe; internasal width 1.2 7-1. 13 in first toe;
internasal width i. 5-1. 66 in second toe; internasal width 3.27-3.17 in third
toe; internasal width 3.45-3.33 in fourth toe; internasal width 3.36-3.0 in
fifth toe.

HABITAT

Its original describer, LeConte (1855, p. 427), gives it as an inhabitant
"in ditches and ponds." Deckert (1915, p. 22) found it "in great numbers
about ditches and bayous." In 191 7 Fowler (p. 39) secured it in a hammock.

In 192 1 (Aug. 5) F. Harper just outside the swamp at Fargo, Georgia,
noted that "After considerable search one of the frogs was located in a thick
clump of sedge, its body half submerged."

In 1922 on June 15 near Chesser Schoolhouse in forenoon, (U. S. Geol.
Sheet, Folkston sheet) "under chips, small boards and logs found 3 Pseudacris
in dry pine barrens, associated with Bufo quercicus. Area beneath boards and
cones more or less dry." On July 2, we "went to night hawk nest area
{Pseudacris nigrita spot). At 6:00 p. m. heard no P. nigrita. Heard one or
two P. ocularis in the grass."

On July 15, 1928, "on high sandy ridge south of Spanish Creek, Folkston,
Georgia. Quercus cateshaei, Quercus cinerea, Gaylussacia dumosa and a very
much serrated poison ivy {Rhus). Miles and I were looking at the Rhus
carefully and gingerly. He saw something jump amongst it. Thought it a
grasshopper but he soon found it to be a frog. It was P. nigrita. The frog
was first on a dead twig of a fallen branch. It would leap amongst dead
leaves, amongst some grass-like grasses and amongst the ivy."

On August 16, 1928, "In making a detour on to the Dixie Highway about
1,2-1.3 miles south of Hilliard, Florida, we came to the road opposite a wet
pine woods. In one little temporary pond in a tussock of grass beside the
small pool was a Pseudacris nigrita calling."

FIRST APPEARANCE

We suspected its presence long before 1921-1922 but not until 192 1 did
we find it. This led us to believe it a very early spring or late winter breeder.
Deckert (1914 and 1915) "found (it) in the winter and early spring." Our
first record of June 15, 1922, is of no consequence in this category. Students
and friends (Dr. A. A. Allen and others) of the author have described from
Florida in February and early spring swamp cricket frogs which must be
this species though slightly rougher than our specimens (see Cope's C. ver-
rucosus). C. S. Brimley received it February 10, 1898, Bay St. Louis, Miss.
O. C. VanHyning has secured for me croaking P. nigrita at Gainesville, Fla. in
February and early spring.

GENERAL HABITS

Metachrosis. Miss Dickerson (1906, p. 157) wrote that the combined
{triseriata-feriarum-nigrita) group was "Changeable from a colour so dark
that it is nearly black, to flesh colour. When light, the colouration may be

200 Frogs of the Okefinokee Swamp

bluish or ash grey, fawn colour, or even salmon or red in tone." This applies
quite well to P. nigrita in its restricted sense except that we have not recorded
any salmon-red individuals yet, though they are possible.

On August 5, 192 1, F. Harper came on one live frog which "was blackish-
olive, whitish below, and had a greenish-white maxillary streak. After a
couple of days of captivity in a can, its black back had changed to olive,
coarsely spotted with black."

Variations in color. Back: Black, speckled with small white warts; middle
of back cinereous with an interrupted stripe of black. — LeConte 1825, Harlan
1825 and 1835.

Back olive brown, with a tinge of yellow, and an interrupted black ver-
tebral line — Holbrook 1842.

Color above dark slate, with more or less of oblong, oval or round some-
times very numerous black spots, which often form stripes. — LeConte 1855.

Gray or greenish-black, with usually three longitudinal rows of black,
light-edged, roundish spots. Occasionally these spots fuse into bands. . . .
A narrow black band starts at the tip of the snout, runs through the eye,
covers the tympanum and ends at the shoulder or beyond — Deckert 1915.

Blackish ash above, with three interrupted black longitudinal bands;
loreal and temporal regions black — Boulenger 1882. Color above leaden, with
three longitudinal rows of darker, light-edged spots, extending one on each
side, and one on the median line. These spots may be united into a band on
one or on both sides or on the middle hne. Cope 1889.

Blackish ash, with three or five darker, sometimes interrupted longitudinal
bands; a black streak passing through the eye above the tympanum — Gun-
ther 1858.

Belly. Beneath granulate whitish. — LeConte 1825.
Beneath whitish granulated. — Holbrook 1842.
Abdomen yellowish-white. — Holbrook 1842.

Beneath whitish, sometimes tinged with yellow with a few black spots
assembled in groups cancellately reticulate. — LeConte 1855.

Beneath white immaculate. — Boulenger 1882.

Inferior surfaces yellowish. — Cope 1889.

The under surfaces are greenish-yellow. — Deckert 191 5.

General habits. To Deckert (191 5, p. 23) this is a shy species, extremely
difficult to catch. Holbrook (1842, Vol. IV, p. 108) holds ''But little is known
of the habits of Cystignathus nigritus; but, from the form of the posterior
extremities and the arrangement of the toes, it would seem to be a land
animal, and closely alhed with the Cystignathus ornatus." In general we
suspect it accords closely in habits with P. triseriata and P. feriarum. From
remarks and observations of O. C. Van Hyning at Gainesville, Florida, this is
quite likely true.

VOICE

"Its call is similar to that of the cricket 'frog,' but much louder, and the
crepitations are slower." (Deckert 1915, p. 23). In 1921 at Fargo, Ga., F.
Harper "During downpours on the afternoon of August 5, . . . began to hear

Pseudacris nigrita 201

several frogs calling in a field grown with grasses and sedges and flooded with
rain. The notes consisted of a shrill, metallic, staccato trill, frequently re-
peated: i-i-i-ik, i-i-i-ik, i-i-i-ik. They were indistinguishable from those of
representatives of this genus in the Athabaska region (septentrionalis) and in
the District of Columbia (feriarum)." (We are not including notes of our
experiences with triseriata, feriarum, etc. forms in New York, Pennsylvania,
Canada, Texas and other places nor are Hay's, Dickerson's or other character-
izations included. Their notes pertain largely to northern forms and they
felt, possibly rightly, that triseriata, feriarum and nigrita were all forms of
P. nigrita. There are those who also would put P. septentrionalis in the
nigrita group).

In 1922 we found it June 15 but not calling. On July 15 we took another
"near where we thought we heard it when we were after Hyla squirella at
night." Then on August 16 near Hilliard one was heard after a very intense
rainy spell of miry detours. To me this form which was finally captured
"sounded like our New York Pseudacris." To F. Harper it sounded like a
metalhc, staccato trill, five notes, ic, ic, ic, ic, ic. All our records in July and
August are clearly sporadic croakings long after breeding and after heavy
rainfall.

MATING

Coloration. (The colors given are from life June 15, 1922, but not from
sexed specimens. Specimens, 17 mm. long).

Back drab gray, smoke gray, darb or grayish olive. From snout down mid-
dle of back is one row of more or less connected dusky drab or deep brownish
drab spots. Stripe along jaw pale rose purple or pale salmon color or cart-
ridge buff with some seafoam green.

The row of continuous spots down back at half way point becomes two
rows of separate spots. On each side is one more row of separate spots. Dark
area from snout through eye, back of eye and extending along the side to
groin. Cross bars of legs same color as spots of back and light thin bars be-
tween the spots of legs like background of the back. Under parts white. Iris
black with brownish vinaceous dots. At times iris has considerable of dass
green.

Structural differences. A male (August 16, 1922) of 25 mm. has the
throat darker than the rest of under parts or rather more yellowish (in spirits) .
On the throat are four longitudinal plaints. Possibly the thumb is somewhat
enlarged.

OVULATION

Of the form P. nigrita, in its restricted sense, we have no observations
except negative evidence. In 192 1 we were within the swamp from April 25
onward but it was not until August we found it. In 1922 we were on an island
near the eastern edge from June 14 to September i or later and found a few
specimens on the mainland, none calling except one in August. The breeding
must have been early and most tadpoles transformed before June 15. Deck-
ert found them active and calling in winter and early spring.

202 Frogs of the Okefinokee Swamp

EGGS

(Not described). Doubtless like those of Hay's (Indiana), Morgan's
(Maryland), Wright's (New York) swamp cricket frog observations and those
of several other observers. We also have Pseudacris eggs from Texas and
tadpoles from several diverse parts but no eggs of P. nigrita as such are on
record.

TADPOLES

(Wherein these differ from Texas, Indiana, Maryland, New York and
Louisiana tadpoles has never been observed or recorded by any South Car-
olina to Mississippi naturalist or visitor to those parts).

LARVAL PERIOD

In the northern regions kindred species take 67 days (Hay's), 60-70 days
(Wright) or about two months as larvae. In 1922 we found none on June
14 on our arrival nor any thereafter. Most of the breeding must have been
over before April.

TRANSFORMATION

In general it must take place before June i. In the North the bulk of
larvae of the related species are transformed by June 15 or by the end of June,
and it must come earlier in Georgia. The northern forms transform at about
9-12 or 13 mm.

GROWTH

Possibly with transformation presumably from 9-12 or 13 mm. the frogs
taken June 15, 1922, which were 17 mm. are one-year-olds. Those taken
July 15 and 18 and August 16, 1922, were 22, 23 and 25 mm. Are they two-
year-olds and does the species breed at two-year-old stage or do some even
enter breeding after one year? We know not.

ENEMIES

This species cannot help being one of the abundant foods of snakes and
numerous foes in early spring through no definite records appear.

AUTUMNAL DISAPPEARANCE

Our latest records are: August records in 1921 and 1922, September 8,
1910, and September 11, 1919.

AFFINITIES

LeConte (1825, p. 282) placed Rana nigrita right after "Rana gryllus
(Savannah cricket)." Harlan (1825, p. 341; 1835, p. 105) repeats LeConte's
"indication" of a fuller description. Dumeril and Bibron (1841, Vol. 8, p.
509) places it in their new genus Acris with the "palmure des pied tres-
courte" as contrasted with "bein developpee" in Acris gryllus.

In his Catalogue of Reptilia and Batrachia Holbrook (1849, Appendix p.
15) still uses Cystignathus.

Pseudacris nigrita 203

LeConte (1855, p. 627) writes "This species has been placed by Dr. Hol-
brook in the genus Cijstignathus, to which it cannot possibly be referred, as it
has the vocal vesicle under the chin and not at the corners of the mouth. M.
Dumeril has called it an Acris. That genus has the toes properly and strongly
webbed."

In 1858 Gunther (p. 97) places it in Fitzinger's genus Pseudacris. Four
years before Baird (1854, p. 60) established Helocaetes for H. feriarum, H.
triseriatus and H. darkii, and Chorophilus for C. nigritus. Chorophilus he
held was "Distinguished from Acris by expansion of apophyses; from Litoria
by the emarginate tongue, less membrane of toes, and more posterior palatine
teeth ; from Hylodes in having a membrane at base of toes, etc." Helocaetes was
different "from Chorophilus in more anterior position of vomerine teeth, etc."

From 1882 (Boulenger) onward until 19 17 (the first edition of the Check
List of Stejneger and Barbour) Chorophilus has remained. Until 1892 C.
triseriatus, C. feriarum, C. nigritus traveled as separate species but at that
date O. P Hay put them all under C. nigritus as subspecies and Miss Dicker-
son followed his treatment. The authors of the Check List, through they
separate them, were not satisfied with the status of the species or forms of
Pseudacris nor is any one today. P. ocularis is distinct. P. ornata is dis-
tinct. Is P. occidentalis a good form, or near if not P. ornata'^. Are P.
nigrita, P. triseriata, P. feriarum and P. septentrionalis all one? We have
worked out one life history in Southern Texas, one in Pennsylvania and one in,
New York. About two more life histories ought to give enough ammunition
to determine this question. We have hastily glanced over the U. S. National
Museum Pseudaais material but more fresh material is needed.

In 1889 Cope said he described another form C. verrucosus from Florida.
Possibly it is a good separable from but Pseudacris in Florida and Texas tend
to become more warty, as do often Acris. Cope held C. nigritus to have
longer hind legs, tubercular upper surface peculiar and swollen gular region
wanting in Northern frog, i.e., C. feriarum.

In 1923 Noble on the basis of a "huge series" (336 specimens) of P.
ocularis collected by Mr. J. A. Weber, some P. occidentalis and comparative
P. septentrionalis material concluded that Pseudacris has no characters
distinguishing it from Hyla and it must be referred to Hyla. He may be
right but we are still following Stejneger and Barbour's Check List (1923) in
retaining Pseudacris until it is carefully and thoroughly worked out. For
example, we have lived with P. ocularis in 1914, 1921, 1922, 1928. It is a
distinct species. We lived with P. ornata in 1925. It is a distinct form from
the P. triseriata, P. nigrita, P. feriarum, P. septentrionalis group. Life
history notes of this complex have been published by Hay, Morgan and others.
We have worked them out in Texas and two different places in New York and
collected tadpoles from New York to Texas. This complex is the crux of the
question, not P. ocularis. Until the puzzling forms are worked out (P
triseriata, P. feriarum, P. septentrionalis, and P. nigrita, P. verricosus) are
carefully studied in situ and in laboratory it is better to keep the genus
Pseudacris. Subsequent investigations may prove realignment desirable.

204 Frogs of the Okefinokee Swamp

BIBLIOGRAPHY

1854 Baird, S. F. Descriptions of New Genera and Species of North American Frogs.
Proc. Acad. Nat. Sci. Phila. Vol. VII, p. 60.

1914 Deckert, R. F. Copeia, Feb. 14, 1914, No. 3.

1915 . Copeia, July 27, 1915, No. 20, pp. 22, 23.

1842 DeKay, James E. Zoology of New York, Part III. Albany, 1842, p. 65.

1841 Dumeril, A. M. C. et G. Bibron. Erpetologie Generale, Vol. VIII, 1841, p. 509.
1917 Fowler, H. W. Copeia, Apr. 24, 1917, No. 43, p. 39 (wrongly labelled p. 27).

1858 Gunther, A. Catalogue of the Batrachia-Salientia in the Collection of the British

Museum, London, 1858.
1825 Harlan, R. Journ. Acad. Nat. Sci. Phila. Vol. V, Part II, Phila., 1827, p. 341.

1835 . Medical and Physical Researches Phila. 1835, p. 105.

1892 Hay, O. P. The Batrachians and Reptiles of the State of Indiana. Indiana Dept.

of Geol. and Nat. Resources. 17th Ann'l, Report. 1891. Indianapolis, 1892, pp.

409-602.

1842 Holbrook, J. E. North American Herpetology, Vol. IV, 1842, pp. 107, 108.

1849 Holbrook, J. E. Catalogue of Reptilia, Amphibia and Fish of Georgia. In White,
G. Statistics of the State of Georgia, Savannah, 1849 (Appendix, p. 15).

1825 LeConte, John. Annals Lyceum of Nat. Hist. N. Y. Vol. V, Part 2, N. Y. 1825,
p. 282.

1856 LeConte, John. Descriptive Catalogue of the Ranina of the United States. Proc.
Acad. Nat. Sciences Vol. VII (1864, 1855) Phila. 1856, p. 427.

1922 Loding, H. P. A Preliminary Catalogue of Alabama Amphibians and Reptiles.
Geol. Survey Ala. Mus. Paper No. 5, Ala. Mus. Nat. History, University Ala-
bama 1922.

1923 Noble, G. K. The Generic and Genetic Relation of Pseudacris, the Swamp Tree
Frogs. Am. Mus. Novitates No. 70, Apr. 23, 1923, pp. 1-6.

1927 Pickens, A. L. Amphibians of Upper South Carolina. Copeia No. 165, Oct. -Dec.
1927 (Dec. 23), p. 109.

Pseudacris occidentalis (Baird and Girard)

(Account written in 1929)

SMOOTH CHORUS FROG

In 1909 in September Professor J. C. Bradley took a frog which we then
identified as Chorophilus occidentalis. Not until 19 12 did we see Pseudacris
aUve in Georgia and Florida first hand in the field and the identification is
uncertain and doubtless incorrect. Cope (1889) pronounces two specimens
taken by W. J. Taylor at Allapaha, Georgia, as C. occidentalis. This locaHty
is in the upper Suwannee drainage. We have not yet critically examined
these two specimens.

Holbrook in his own region (South Carolina) recognized in 1842 three
species: Cystignathus ornatus, Cystignathus nigrita and Hylodes ocularis — the
three distinct groups of today. In 1849 he gives the same three species.
LeConte (1855), another resident of these parts (in Georgia at Riceboro
whence Cope's C. occidentalis material largely came) gives three species (of
the three types) : Chorophilus nigrita, Chorophilus ornatus and Hylodes ocu-
laris. Apparently LeConte did not honor or recognize C. occidentalis at Rice-
boro, Georgia. Cope when distinguishing C. occidentalis makes it the eastern
representative of C. ornatus which he has from Texas.

But Boulenger in 1882 does not help matters by recognizing both C.
ornatus and C. copii (C. occidentalis) in the same region. Sometimes it is hard
to believe that both Holbrook and LeConte are so lacking in taxonomic sense
and that old museum material from LeConte can be acclaimed new or C.
occidentalis by Cope and Boulenger. Rather what these cataloguers have

Pseudacris occidentalis 205

called C. occidentalis in the Southeast is doubtless what the resident naturalists,
Holbrook and LeConte, called C . ornatus. The only collector and resident in
recent years who has Hved in the Southeast long enough to collect and observe
three forms of Pseudacris is R. F. Deckert (1913-1915). In 1913 he found C.
nigritus, C. ocularis and C. occidentalis, the latter "plentiful, in scattered
companies." In 1915 he gave us the only detailed first hand account since the
description of C occidentalis.

His whole account is worthy of repetition. His descriptions and accounts
read like our experiences with P. ornatus in Helotes and Beeville, Texas, in
1925. His notes are:

"Chorophilus occidentalis, B. & G. is a beautiful species, rarely seen except
in early spring. At this time, however, it occurs in great numbers in the large,
shallow bayous, where it stands almost straight upright, on some twigs or
weeds, with its large throat pouch distended, giving vent to its piercing call.
This call is very loud, similar in pitch to that of Hyla pickeringi, but much
shorter, and at a distance sounds like the ring of a steel chisel, when struck
with a hammer.

"This is the largest species of the genus Chorophilus, attaining a length of
I 1/2 inches, from snout to vent. It is stout of body, the head is pointed, the
arms and legs rather short and stout. The toes are very shghtly webbed, the
disks on fingers and toes scarcely noticeable. The subarticular tubercles are
very prominent.

"The structure of this species indicates terrestrial, possibly subterraneous
habits. I have dug specimens out of the sweet potato hills in my garden.

"The smooth, shiny upper surfaces are rich reddish brown or fawn colored,
like specimens of our northern Wood frog. There is a black band on each side,
beginning at the nostril, running through the eye, over the tympanum to the
shoulder. The edge of the upper jaw is silvery white. The groin is rich yellow
or orange, with large elongate or round black spots. The undersides are white,
faintly spotted with brown at the throat, flesh colored posteriorly. On the
back there may be two broad, dark brown parallel bands. The arms and legs
are banded more or less distinctly with brown." This was the only one of
Deckert's Jacksonville Anuran species we missed. We did not arrive early
enough to find it in activity.

The P. occidentalis- P. ornata difficulty can be easily settled with new
material taken from South Carolina to Texas. Pseudacris does not have nine
species in U. S. A. Too many personal equations enter the problem with
scanty material. And Litoria occidentalis Baird and Girard may be a Riceboro
(Georgia) form as Cope believed but is it not C. ornatus of LeConte, its col-
lector, and a familiar student of C. ornatus at Riceboro, Georgia? He never
said Riceboro, Ga., had both C. ornatus and C. occidentalis. To populate
South Carolina to Florida with 5 species of Pseudacris is overcrowding and the
placing of the so-called San Francisco Litoria occidentalis (1853) in the South-
east was not apparently honored by LeConte two and one-half years
later (December 1855,) in his Descriptive Catalogue of the Ranina of the
United States. LeConte himself collected Baird & Girard's L. occidentalis

2o6 Frogs of the Okefinokee Swamp

types in 1850, yet five years later he omits, forgets or disregards them. Still
in fairness it must be said he does not mention the other three species of that
paper: Bufo halophila, Hyla regilla or Rana LeContei.

In the Amphibians and Reptiles of Arkansas Hurter and Strecker record
Chorophilus occidentalis from "Hot Springs (Combs)". Strecker knows
Pseudacris ornatus and has contributed as much or more to its habits than any
one living but the Pseudacris occidentalis is doubtful.

In December 1927 Mrs. Wright and I superficially examined the Pseudacris
ornata, P. occidentalis material of the U. S. National Museum. The material
of the latter species is generally rather old. A few tentative notes we made
were that No. 5905 C. occidentalis of Cope's identifications is P. ornata.
Some specimens from Florida labelled P. occidentalis are P. ocularis without
vitta. Some P. occidentalis collected by Hurter in Mississippi look to be P.
ornatus. Another specimen collected by J. Hurter from Waco, Texas, was
labelled Lithodytes latrans and proved to be Pseudacris ornata. This is a little
of the apparent mixed condition of Pseudacris with little good comparable
museum material. No hope comes except from considerable collecting and
field familiarity with the forms themselves. We have not yet begun a critical
study of P. ornata-P. occidentalis nor the P. nigrita complex but immediately
plan such a study.

Since the above was written we remembered that C. S. Brimley (1910, p.
11) secured "Chorophilus occidentalis from Bay St. Louis, Mississippi, Feb-
ruary and April 1898, 5." and Chorophilus ornatus "Hastings, Florida, June
1901, i; Green Cove Springs, Florida, July, 1898, 5." He is the only person
to receive each from the Southeast.

Pseudacris ocularis (Holbrook)

(PL I, Fig. 7; IV, Fig. 6; V, Fig. 6; VI, Fig. 4; X, Fig. 8; XII, Fig. 8; XV, Fig. 8; XVII;
XXIV, Text Fig. I, 6)

COMMON NAMES
Swamp Tree Frog. Swamp Cricket Frog. Little Chorus Frog. Tree Frog.

RANGE

Check list. "Type Locality: South Carolina and Georgia. Range: South-
eastern United States"— Stejneger & Barbour Check List (1923, P- 28).

Supplemental records (or detailed account of records) . Outside of the Oke-
finokee parties from 1912-1922, the one student of Amphibia who has actually
known this species first hand and best is R. F. Deckert. Subsequently, Mr.
Jay A. Weber and others secured material for Dr. Noble and more recently
Mr. Vernon Bailey has taken it in Florida for the U. S. National Museum.
In 191 1 and 1912 R. F. Deckert (Copeia, 1914, p. 3) found it "abundant and
everywhere" ten miles south of Jacksonville. In 191 5 the same author (191 S>
p. 23) reports them "very numerous near Jacksonville" .... In 1920 he

Plate XXIV

Black chorus frog; (Pseudacris ocularis)

1. Saddlebag pond, Tiail Ridge, Folkston, Ga. July 9, 1922.

2. Male croaking on trunk of black gum, Billy Id., Ga. May 20, 1921. Flashlight.

3. Male croaking on bush, Chesser Id., Ga. July 26, 1922. Flashlight.

4. 5. Mature tadpoles, Saddlebag pond. Trail Ridge, Folkston, Ga. June 30,

1922. X 1.0.

6. Tadpole with two legs. Saddlebag ])ond. Trail Ridge, Folkston, Ga. June 30,

1922. X 1.0.

7. Tadpole with four legs, Trail Ridge, Folkston, Ga. June 30, 1922. x i.o.

8. Tadpole with short tail, Chesser Id., Ga. July 6, 1922. x i.o.

9. Transformed frog, Saddlebag pond. Trail Ridge, Folkston, Ga. June 30, 1922.

X I.o.

10. Male croaking on twig four feet above water, Billy Id., Ga. June 24, 1921.

Flashlight.

1 1 . Habitat of Pseudacris ocularis, in gi-ass and small bushes of foreground. Trees

and bushes of Siren Run in background, Billy Id., Ga. May 27, 1921.

12. Male croaking on grass blade, close to the water, Billy Id., Ga. July 6, 1921.

Flashlight.

t f ^ f J^ it

Plate XXIV

Black chorus froo- (Pseudncris ocularis)

1. Saddlebag i)()nd, Trail Ridge. Folkstcm, Ga. July 9, 1922.

2. Male croaking on trunk of black gum, Billy Id., C5a. May 20. 192 1. Flashlight.

3. Male croaking on bush, Chesser Id., Ga. .July 26, 1922. Flashlight.

4. 5. Mature tadpoles. Saddlebag ]i()nd. Trail Ritlge, Folkston. Ga. .Tune 30,

1922. X I.e.

6. Tadpole with two legs. Saddleliag jwnd. Trail Ridge, Folkston. Ga. June 30,

1922. X i.o.

7. Tadpole with four legs, Trail Ridge, Folkston, Ga. June 30, 1922. x i.o.

8. Tadpole with short tail, Chesser Id.. Ga. July 6. 1922. x 1.0.

9. Transformed frog. Saddlebag pond. Trail Ridge, Folkston, Ga. June 30, 1922.

X I.o.

10. Male croaking on twig four feet above water, Billy Id.. Ga. June 24, 1921.

Flashlight.

1 1 . Habitat of Psciidocris ocularis, in grass and small bushes of foreground. Trees

and bushes of Siren Run in background, Billy Id., Ga. May 27, 1921.

12. Male croaking on grass blade, close to the water, Billy Id., Ga. July 6, 1921.

Flashlight.

f f I f J^ 4^

Pseudacris ocularis 207

(Copeia, 1921, p. 22) finds it quite numerous in Dade County. In May, 1922,
he recorded (Copeia, 1922, p. 88) it again in Dade County. In 1923 Noble
(1923, p. 3) had specimens from Lee Co., Florida, Arlington, Florida, near
Jacksonville, Florida, Hampton Co., S. C. and Wilmington, S. C. Noble did
not intend Wilmington, South Carolina, for G. S. Myers (1924, p. 59) re-
cords "I know of no other specimen being recorded from the State (N. C).
It is the one mentioned by Noble (Am. Mus. Novitates, 70, p. 3) from Wil-
mington." Wright (1926, p. 82) gives Pseudacris ocularis as a Sabalian species
from South Carohna to Florida. Inasmuch as Wilmington, N. C. is Sabalian
country it is not surprising to find it recorded there. Its range seems to be
from N. C. to Dade and Lee Counties in Southern Florida. The last record is
Brimley's (1926, p. 82). He records the "Little Chorus Frog {Pseudacris
ocularis) from "(Pitt, Craven, New Hanover)" counties.

The earlier material is somewhat as follows: Holbrook (1828 or 1842) has
specimens of Hylodes ocularis from S. C. or Georgia. Several years later he
(1849, App. p. 15) records this species from Georgia. LeConte (1855, p. 429)
says Hyla ocularis "inhabits Georgia." In 1875 Cope (1875, p. 30) dubs it
Chorophilus angulatus. Boulenger (1882, pp. 333, 334) probably had no
material at hand. Cope (1887, pp. 348-349) had not Girard's specimens from
Charleston, S. C. and relied on Baird's manuscript notes. Until 1911-1928
it was not at all common in collections.

Local Okefinokee records. In 191 2 we wrote "In all we have ten specimens
from the swamp, one taken May 28, 19 12, one May 30- July 15, 191 2, two
June 21-22, 1912, three June 24, 1912, two July 1-15, 1912, and one after
July IS, 1912."

In 1 92 1 on Sunday, July 3, we took a handcar trip to Honey and Black
Jack Islands. It was a misty day at first, then sunny, and rained all the
afternoon. The temperature was from 'ji°-'j6°. On the sphagnaceous strand
of Honey Island heard these creatures, near Black Jack Island, and on
Honey Island in mid afternoon, in Cypress bay between the Pocket and Jones
Island, and all along the trestle between Jones Island and Billy's Island.
Later and earlier on Billy's Island we found them in several locations. On
Floyd's Island and Chesser's Island we also heard and saw them.

In 1922 we secured them on Chesser Island, near old Suwannee Canal,
along road to Folkston, in ponds near Trader's Hill, near an overgrown
sphagnum bog outside the swamp along the road from Folkston, Ga., to
Jacksonville, Fla.

GENERAL APPEARANCE

Holbrook speaks "of this beautiful little animal, as the smallest of the
frog kind with which I am acquanited . . . ." Its general characters he gives
as follows: "Head long; snout pointed; upper jaw white, which colour is con-
tinued to the anterior extremity, above this a black band runs over the shoulder
and terminates on the flanks; body short, chestnut-brown or bronzed; ab-
domen yellowish- white." DeKay (1842, Part III) gives "Hylodes ocularis.
(Holbrook, Vol. 3, pi. 14; and Vol. 4, pi. 35, 2d Ed.) Very small; Reddish
brown ; a black band from near the end of the snout, runs through the eyes and

2o8 Frogs of the Okefinokee Swamp

along the flanks. Length three-quarters of an inch. South Carolina and
Georgia." LeConte, 1855, calls it "The smallest of all known Ranina. From
the small size of this and the preceding species, the web between the third and
fourth toe is not very perceptible." Boulenger (1882) calls it "extremely
small." Cope (1887, p. 348) says that "its most striking features will be
found in the large size of the eyes, the acuteness of the head, the small tongue,
etc. The tibia is longer than in any of the genus." To Deckert (1915, p. 23)
"This is the smallest of the North American frogs, rarely exceeding 2/8 inch
from snout to vent. The head is pointed, the eyes large, the limbs and body
slender. The disks on the fingers and toes are small but distinct. The skin
of the back is covered with very fine warts, that of the belly finely granulated.
Evidently Miss Dickerson (1906, p. 162) did not have the species at hand. Of
it she writes "This species is said to be the smallest among North American
Hylas. Its unusual characteristics seem to be the pointed paw, the long legs,
and the light line along the outer edge of the tibia."

C. S. Brimley's (1926, p. 82) revised key summarizes or distinguishes the
little chorus frog as having "snout truncate in side view, adult about two-thirds
of an inch in the length of head and body."

These are the brownies of frogdom in the United States. They are usually
uniform gray, brown or greenish on the back, with a vitta from eye backward
as a stripe of variable length. The snout is pointed, projecting. They cer-
tainly at first acquaintance impress one as the possible young of some Hyh.
We thus puzzled and most authors who have seen them alive have gone
through the same stage, particularly if they are green like so many trans-
formed Hylas.

COLORATION OF SPIRIT SPECIMENS (191 2)

The original color notes of Holbrook which have been constantly repeated
are: "the pupil black, and the iris grey, with a tinge of red. The upper lip is
white, and this color is continued in a line to the axilla; above this is a black
blotch which begins very narrow and becomes broad under the orbit, passes
over the shoulder, and there terminates. The chin and throat is yellowish-
white. The body — uniform chestnut-brown or bronze-coloured above and
yellowish-white below. The anterior extremities — brown above and clouded
white below; — The posterior extremities — reddish brown above, barred with
dusky, and obscure white below; . . . ." LeConte (1855, p. 429) describes
it as "Above brown or bronzed or silvery grey, very finely specked with dusky
or darker, a tolerably wide band of black proceeds from the tip of the nose to
the middle or beyond the middle of the sides, this is bordered beneath with
white. Chin and under side of the thighs speckled with black. Legs speckled
like the back and more or less spotted and barred with dusky, fingers and toes
all furnished with small disks."

Cope in 1887 gives its general color as dark chestnut above, gives a median
dorsal stripe which bifurcates behind and says it can be distinguished from
other Pseudacris by its chestnut color Deckert (19 15, p. 23) who has seen
many characterizes it as follows: "The color is yellow, reddish brown or

Pseudacris ocularis

209

chestnut. The upper jaw is margined with white. There are three longitud-
inal dark brown dorsal bands, one from tip of the snout to above the vent,
usually bifurcate posteriorly, and one on each side of this, starting behind the
eye. Underneath, it is pale yellow. The males throat is dark brown, and can
be distended to the size of a large pea. The arms and legs are distinctly cross-
banded." Deckert thus is one of the first to describe the male and adds yellow
to the general color.

Weber (Noble 1923, p. 4) finds them green in life. Noble (1923, p. 5)
finds the lighter phase of more frequent occurrence and without mid-dorsal
fine of dark brown while the less frequent type "dark phase has three longi-
tudinal stripes of dark brown on the back, very similar in form to P. triser-
iata.'^

Color has been too strongly emphasized in this group. From just chestnut
or bronze in Holbrook we have had silver grey, yellow and finally green added
to general descriptions of its color. Cope emphasized the chestnut as dis-
tinctive. All the above colors may be recorded, though the author has seen
fewer green specimens than of any other phase. The Florida collectors (see
Deckert, Noble) often mention the mid-dorsal stripe but we have very few
with it. Yet a recent collection from Florida made by Mr. Gerritt S. Miller
have the general type without dorsal stripes and only the lateral stripes as in
Holbrook's figure, but examples appear with mid-dorsal stripe or supplemental
ones on either side of it.

Some rough notes we made on this collection of 8 males made by G. S.
Miller Jr. and C. R. Aschemeier are: "The vocal sac of the male, even when
collapsed, sometimes appears to cover almost half of venter of body, the rear
of it may reach caudad to the pectoral region (fine connecting one arm in-
sertion with the other).

U.S.N.M. No. 71038. Has no dorsal stripe but has dark dots on back and
tibia sometimes assembhng in somewhat larger ill-defined spots. The
entire vertical parts with dots like those of the back, but much wider
spaced. Sometimes this dark punctate arrangement becomes very pro-
nounced.
U.S.N.M. No. 71036. No dorsal stripes.

U.S.N.M. No. 71039. No dorsal stripe. If anything, these males from
Gainesville seem to be more tuberculate on the venter than do those of the
Okefinokee.
U.S.N.M. No. 71040. Has the venter from somewhat caudad of pectoral line
more tuberculate than almost any other hyhd in the U.S.A., but not on
hind legs. Hind legs big proportionally. Color fawn brown on back. No
dorsal stripes.
U.S.N.M. No. 71041. No dorsal stripes.
U.S.N.M. No. 71034. No dorsal stripes.
U.S.N.M. No. 71035. Darker back. No dorsal stripes.
U.S.N.M. No. 71037. Has suggestion of indistinct stripe down mid dorsum
and on either side of it is a dorsolateral stripe toward the rear. The eye
vittae go just back of axillae. There is a prominent dark stripe on the front

2IO Frogs of the Okefinokee Swamp

of the tibia. This coloration suggests the typical Pseudacris coloration,
there is a dark spot on the top of the snout. None, however, have a tri-
angle between the eyes.

STRUCTURAL CHARACTERS (WRITTEN IN 191 2)

Upper parts smooth; the under parts also smooth, except for the ventral
parts back of the ventral fold, the upper breast and belly and under side of
the thigh granulate; nostril lateral and much nearer the tip than the eye;
snout pointed and projecting; lower jaw narrow; cleft of the mouth deep, to
the vertical of the tympanum; in most of the specimens a prominent breast
fold present ; in all, tongue entire and in most broad ; in the smaller forms where
lower jaw is sometimes narrower, tongue is sometimes narrow; tympanum
.5-3 of the eye; disks moderate on the feet; disks more prominent on the
fingers; first finger shorter than the second; second and fourth fingers about
equal; third finger longest; subarticular tubercle small; a metacarpal tubercle
present; fingers free; metatarsal tubercles external and internal small; sub-
articular plantar tubercles indistinct ; fourth toe much longer than 3rd or 5th
which are about equal; 2nd sHghtly larger than the first; toes hardly half
webbed; in some of the larger specimens there is present an indistinct fold
from eye over tympanum to the arm insertion.

MEASUREMENTS

(1912-1914)
In 1912 the measurements of the ten individuals were "from 12-15. 5 mm.,
average 12.5 mm.; the head is 3.5-6 mm., average 4.5 mm., more than the
width of the head (like Boulenger's C. copii); head contained in the length
2.3-3.4 times, average 2.75, mode 2.9; the width of the head, 3-4.5 mm., av-
erage 3.75 mm., 3-4.3 times in the length of the body; about the same or
narrower than Cope's 3-3.5 for C. occidentalis; snout 1.5-2.75 mm., average
1.36, from 1-2 times the eye rarely i times and usually about 1.5 times the
eye; the eye is from 1-2 mm.; interorbital distance which is .5-2.25 mm.,
average 1.5 mm., equal to or greater than the eyelid which is .75-1.5 mm.,
average i mm. The femur is 5.25-8 mm., average 6.5 usually less than the
tibia, occasionally equals tibia; tibia 5.75-8 mm., average 6.8 mm., usually
continued 1.75-2 times in the length of the body; tarsus 3.5-5.25 mm., average
4.4 more than the width of the head and usually equal to the length of the
head; rest of the leg 4-6.25 mm., average 4.9 mm., equal to or slightly more
than the tarsus; anterior limb from axilla 5.25-7.75 mm., average 6.4 mm.,
about equal to femur; posterior limb 19-23.5 mm., average 20.4 mm., usually
1. 5-1. 6 times the length of the body; usually the heel of the hind limb reaches
snout, occasionally half way between the eye and the tip of the snout.

(Recent Material)
Head to angle of mouth i.o (17.5 mm. 9) in width of head; head to
rear of tympanum .91 in width of head; head to angle of mouth 3.5 in length
of body; head to rear of tympanum 3.18 in length of body. Snout, 0.5

Pseudacris ocularis 211

in first finger; snout, .75 in fourth finger; snout, .66 in first toe; eye, 1.8 in
snout; eye, .5 in tympanum; eye, .85 in first finger; tympanum, 5.0 in inter-
tympanic width; tympanum, 3.3 in snout; internasal width, .5 in upper eye-
lid width; interorbital width, .5 in upper eyehd width; interorbital width, i.o
in internasal width; interorbital width 2.25 in intertympanic width;

Forelimb: Forelimb, 2.5 in length of body; forelimb, t,.^ in hind limb; first
finger, 2.0 in third finger; second finger 1.5 in third finger; second finger, .75 in
third finger; third finger, .83 in second toe; fourth finger, 1.33 in third finger;
fourth finger, .90 in first toe; internasal width, .75 in first finger; internasal
width, 1.0 in second finger; internasal width, 1.5 in third finger; internasal
width, I.I 25 in fourth finger;

Hindlimb: length, 1.3 in hind hmb; tibia, 2.2 in length; tibia, 2.9 in hind
limb; tibia, .875 in forehmb; tibia, .85 in hind foot; first toe, 1.25 in second
toe; first toe, 2.0 in third toe; first toe, 2.75 in fourth toe; first toe, 2.1 in fifth
toe; second toe, 1.6 in third toe; second toe, 2.2 in fourth toe; second toe, 1.68
in fifth toe; third toe, 1.375 in fourth toe; third toe, 1.05 in fifth toe; fourth
toe, 1.23 in hind foot; fourth toe, 1.4 in tibia; fourth toe, 1.2 in fore limb;
fifth toe, 1.3 in fourth toe; internasal width, i.o in first toe; internasal width,
1.25 in second toe; internasal width, 2.0 in third toe; internasal width, 2.75 in
fourth toe; internasal width, 2.1 in fifth toe.

HABITAT

Holbrook (1842, p. 138) refers to this species as "choosing the same damp
places for its abode (as Acris gryllus) and is not infrequently found on the
leaves of such low shrubs as inhabit the same localities, as the myrtle (Myrica
cerifera); . . . ." "... this animal is not infrequently found on low bushes,
or leaps upon them when pursued, . . . ." .... "They are all found near
water, or in darkest places, like to Ranoida, but are never observed sitting half
immersed in it, though they will conceal themselves by diving to the bottom
when pursued."

Deckert (191 5, p. 23) writes "During the spring months, they abound on
the marginal vegetation of the 'bayous," . . . ." In 1922 he (1922, p. 88)
reports them "heard at Homestead and in rain-ditches along the road to a little
beyond Royal Palm Hammock," in Miami district. Mr. Jay A. Weber (Noble
1923, pp. 4, 5) gives his experiences with them as follows: "The tiny frogs
were a brilliant light green in life, similar to, but even brighter than H.
cinerea. They were observed most abundantly at Rocky Lake. A narrow
belt of reeds along the water's edge had been tramped down by cattle, and it
was here the frogs in association with Acris were found. They were extremely
agile and I found it impossible to get near enough to catch them by hand.
As one approached them they would hop among the crushed, but still living
reeds, where their green coloration caused them to disappear immediately.
Never did they seek a hiding place in the holes or shallow puddles in the
manner of Acris. The deep water of the lake was full of fish and I noticed
that neither Acris nor this species ever hopped into the deep water. At last,
after constructing a club from the base of a cat-tail stalk, I succeeded in

212 Frogs of the Okefinokee Swamp

approaching close enough to secure specimens. The species is the rarest of
the riparian frogs in the region that I visited and its extreme agiUty and pro-
tective coloration make it the hardest, by far, to catch."

In 191 2 we have these notes of this species in Okefinokee Swamp. "These
forms were taken generally in rather moist situations. One of the special
places was a narrow marshy crossway between two islands with associated
killifishes and young Centrarchidae. On May 28, 191 2, at the end of the
lumber railroad and near Mixon's Ferry we found some Chorophilus hopping
about in the vegetation with oak toads as associates. On May 10 on Honey
Island Prairie, a different sort of place, there were some small frogs which we
considered Chorophilus but of them we made no collections. The bulk of
the specimens were secured in some small ponds on Billy's Island or in the
damp dark localities among the heath cover."

On May 16, 1921, ''picked up Pseudacris ocularis in the grass, in amongst
pipewort (Eriocaulon)." The next day on Billy's Island we went out to
Crosby Pond. In the grass, sedge and Eriocaulon cover, also some heaths,
took Pseudacris ocularis.'^ The following day, May 18, "at 4:40 p.m. went
to Crosby Pond. Pseudacris ocularis may start up from sphagnum edge of
Cypress Pond where sphagnum is no more than an inch above solid ground
and in the piney wood's side outside the gum border of a pond. May start
this form up from the ground. It will leap i-i 1/2 feet at times. Laid on
Eriocaulon, small Sarracenia minor, sedge and saw palmettos. Took a dozen
of them." On June 8 "we went to Siren Run which is getting very dry. In
one moist place amongst pipe worts {Eriocaulon) found a few Pseudacris
ocularis. ^^

On June 25 we found them common in one pond "Many in bamboo
(Smilax) vines, on bushes some 4-5 feet up. Others on level of water. All
the way from open pond through fringe of gum trees to the outer edge of the
pond, in amongst fallen pine tops, in amongst trash, where water is shallow
and covered with drift particles, etc."

On July 15, 192 1, on a trip to Chesser Island we found them abundant on
the outer edges of cypress ponds outside the swamp, on the outer edge of the
marginal cypress bay. Most of them we "caught in the grass near the edge
of ponds."

In 1922 we found them from Trader's Hill to Chesser Island in similar
habitats to those of 192 1, e.g., on June 17 "in grassy and sedgy area we found
several on the edge of pine barrens on outer edge of cypress bay or branch and
also one mile south of Canal near swamp's edge."

FIRST APPEARANCE

At the very first of our 19 12 trip we found them (May 28). In 192 1 we
entered the swamp the last week in April, one month earlier than above. At
once the boys brought in 3 frogs (April 24) and they were females, which
implies the species was well out. No doubt resident naturialists will find earlier
dates of appearance for this species as for all frogs of the Okefinokee. The
earliest record I can find is Deckert's (192 1, p. 22) note for Miami region.

Pseudacris ocularis 213

"This is the smallest of North American Salientia and was found to be quite
numerous in a dried-up ditch with black muck bottom at Little River, Feb.
23. A dozen specimens were taken and liberated later on at 19th Street,
Miami, in a small hammock. It is the only member of the genus observed
here so far." So, too, G. S. Miller Jr. and C. R. Aschemeier took adult males
(U.S.N.M. 71034-41) at Gainesville, Florida, April 3, 1926, and 0. C. Van
Hyning has taken it from February onward at the same place.

GENERAL HABITS

Metachrosis. On May 21, 192 1, "It is curious that I put all sixteen frogs
in a pan and put it in the desk beside two books. All those on the north side
were of the reddish brown phase. Those on the south side of a gray phase.
The first were beside the deeper reddish binding of Jordan — Evermann's
Fishes of North America and the gray ones near Smith's Fishes of N. C., redd-
ish lighter manuscript holders or covers." On the evening of May 21, 192 1,
during a great chorus of this species they impressed us as much faded out at
night.

Variation in color. Others have recorded individuals: chestnut brown,
bronzed (Holbrook) ; brown, bronze or silvery grey (LeConte) ; dark chestnut
(Cope); yellow, reddish brown, chestnut (Deckert); green (Weber).

In 191 2 we secured the following colors: gray, greenish, light yellowish
olive, dull citrine, chestnut bay. In 192 1 we took individuals: brown, reddish
brown, gray, amber brown, hazel, tawny, buckthorn brown, light grayish
olive, light oUve gray, deep olive gray, cinnamon.

On May 17 we had diverse colorings in a lot of sixteen in one pan. "Some
were tawny, hazel or amber brown on upper surfaces, others buckthorn
brown. One is light grayish olive or light olive gray; another deep olive gray.
The stripe on the upper jaw was almost cream, chalcedony yellow or glass
green. Belly chamois, cream buff, or calomel buff. Iris, cinnamon."

General habits. Holbrook, its describer (1842, p. 138) says this species "is
closely allied to the Hylodes grijllus in its habits, choosing the same damp
places for its abode, . . . ." Under this species he notes "The members of this
genus, 1 ke those of the last, are all diurnal in their habits, or seek their food
by daylight. They all delight in the heat of the sun, and the brighter its rays,
the more merry and noisy are they."

In 192 1 we first met them in numbers May 16 on Billy's Island "In the
grassy pipewort cover they, when disturbed, could jump i-i3^ feet away
like an .4cm. The Pseudacris ocularis would jump into small bushes.- Were
very hard to catch. Of six I saw I caught only two. One I thought I had
killed. His tongue hung out yet he jumped. The jump apparently remains
to the last. The adult Acris will leap into the bushes 3^-i foot high. Are
these last year's Acris? This is only a guess, a possibihty." We soon satisfied
ourselves they were different. On May 21 we "found Pseudacris ocularis
hopping over water like a young bullfrog, usually over a dense mat of grass
in water."

214

Frogs of the Okefinokee Swamp

VOICE

We took these creatures as early as 1 9 1 2 but have no clear recollection or
notes to indicate we distinguished their notes in that year. Apparently the
first person to recognize their notes was R. F. Deckert (19 15, pp. 23, 24) who
reports "males giving vent to their faint calls, which sound more Hke the
chirping of insects than the calls of frogs."

Our first acquaintance with a chorus came the evening of May 20, 1921.
My journal reads thus: "Then went down to the pond east of the negro quar-
ters and it sounded as if bedlam had broken loose. I heard a cricket-like note
everywhere and concluded it was Pseudacris ocularis. Have heard isolated
calls of this note during the day and at night for several days, but it was
dismissed as a possible insect yet my identification has not satisfied me. Later
the boys came. I had found Pseudacris ocularis to be the author of the cricket-
like note. One frog was on a grassy mat; one on a log; another calHng from a
pine bush at edge of water; another on the side of the bole of a tree. They do
not Hke the electric spotUght. One on pine bush after a time worked up into
the leaves or crawled up the branch. We flashed one on the side of a black
gum. Its throat pouch was transparent and we could see through it and
discern the bark behind. This one, one of the boys discovered without flash-
light, merely by the aid of moonlight. Many are in the grass and are hard to
see. We timed the croaks. One gave 49, 52, 55, 58 cheeps in a minute;
another 45, 48, 50, 52, 54, 56, 58, 59, 61, 2, 5, 7 calls a minute; another 30,
32, 34, 36, 38 a minute. We found that they averaged in general about 32
calls a minute with silent intervals averaging one second to one and seven-
eighths or two seconds. In some cases as few as 20 per minute were recorded.
Some would be regular; some irregular. Its note is penetrating and can be
heard 150-200 feet away. It is high pitched, insect-like. In some respects
it is little like the notes of other Pseudacris, yet in other respects it reminds one
of the other species of the triseriata-feriarum-nigrita group. It is an amusing
little creature as it squeezes its slender body and throws out its large sac one-
half the size of the body. It may be perched obhquely on the side of a bole
of a tree, lengthwise of brush clump or more often cross-wise of a branch. It
may not be on a bush at all when calHng. However, one characterizes it be
it as a chirp, trill, cheep, squeak, or as high, shrill, insect-hke, it surely is a
loud piercing call for so little a mite of frog flesh."

In late May, 192 1, when following Pseudacris ocularis vigorously we
concluded from our experiences with it that above a certain required minimum
temperature plays less role in croaking of frogs than humidity. When the
temperature at night became the warmest they might not call if the air had
been dry at that period.

"On July 3, 192 1 we made a trip by lever car to Honey Island, Honey Island
Prairie, and almost to Black Jack Island. Railroad to within i mile of Black
Jack. It rained at lunch and afterwards. On our return we heard Pseud-
acris all over the prairie and in sphagnum of wet thickets especially. On
Honey Island we stopped and caught a few Pseudacris. All along trestle be-

Pseudacris ocularis 215

tween Jones and Billy's Island we stopped and caught a few Pseudacris call-
ing. Never heard such a frog din. Same on Jones Island. Later on Billy's
Island no end of Pseudacris in the hammock near Newt Pond."

Our journal notes of calhng are:

1921
April 25. Two or three frogs brought into camp.
May 16-18. Common in Cypress Pond edge.

" 20. In Chorophihis (Pseudacris) Pond at 4 p.m. no end of Pseudacris calling

by day. In chorus in evening.
" 21 . No end of Chorophihis calling at 8 p.m. Air 70°. Were calling vigor-
ously in afternoon.
" 22. Several heard in mid afternoon. Hear them at night at Newt Pond.
" 24. In moist pine barrens near edge of cypress bay heard two at 4 p.m.,

bright sun. Tonight at 10 p.m., air 77°, a few calling.
" 26. A few Pseudacris calling in Chorophihis Pond and other places.
" 27. Temperature up to 91°. Few calling in mid afternoon. Several heard

at night.
" 31 . Hear one or two Pseudacris ocularis at 11:30 p.m.
June 2 . One calling, several seen this day.

" 3. No Pseudacris calling at Chorophilus tonight, 75°. One heard in

hammock tonight back of camp.
" 4. Heard none cahing except one in grassy field near camp.
" 5. In evening went to Long Pond trestle. Heard plenty of Pseudacris

ocidaris, air 71°.
" 14. Heard one or two Pseudacris ocularis about 8 a.m., Floyd's Island.
" 21. Heard one Pseudacris at Bufo Pond at 10 a.m.
" 2^ . Heard a few.

" 24. After rain common at Chorophihis Pond.
" 25 . Common, plenty at Newt Pond. Several calling at Chorophilus Pond.

Hear one at 10:30 a.m.. Long Pond.
" 28. One heard on Jones Island.
" 29. Several heard early in morning about 5.
" 30. Several heard in outer edge of pond, not middle, 3 p.m.
July I . Quite a few calling by Indian Mound ponds.

" 3 . Heard in Honey Island Strand near Black Jack Island and on Honey

Island all in early afternoon. Later in the Pocket. Commonly heard

between Jones Island and Billy's Island.
" 4. Plenty at Long Pond.
" 6. Pseudacm heard 75-80 yards away.
" 7 . Photographed croakers tonight and last night.
" II . A few heard along Suwannee Canal at 6 p.m.
" 12. Common, heard 1-3 p.m. along canal.
" 15. Saw and heard plenty of Pseudacris on Chesser Island.
" 15. Hear Psewdacns around Bay Pond. We saw and heard plenty on the

mainland and on the Island.
" 16. Immense chorus but hard to hear because of choruses of Hyla gratiosa,

Bufo quercicus, etc. Much laying.
" 17. There are fewer calling tonight, yet they are abundant.
" 18. One or two heard Chesser Island.
" 27. Cloudly. This afternoon, Billy Island, heard a few Psewdacns ocM^om.

A few in the woods. One calling near camp.
" 28. Several heard in piney woods. Rainy. In middle of island where some

gallberry bushes (Ilex glabra) with temperature water are, hear

Pseudacris. Strong chorus tonight.

2i6 Frogs of the Okefinokee Swamp

" 29. Black Jack Island. Down pour near our camp in temporary pool. One
of the greatest choruses of Chorophilus ocularis I ever heard.
Aug. 9 . One or more heard along St. Mary's river.
" II. Many heard along River.
" 17. Several heard at Camp Pinckney, Ga.

1922
June 14 . A few were calling in Coat Bet Pond, Chesser Island at 8-9 and possibly
later.
" 19. Heard a few in one or two places on our return from Folkston.
" 21. Heard one or two beside path from landing into Chesser Isknd.
" 23 . A few calling near the ponds i mile south of Trader's Hill, Ga.
" 26. Heard some in cypress around an open cypress pond.
" 27. Heard a few near Anna's Pond, Starling Branch, Colerain and several
other places.
July 2. 8:30 P.M.-12 midnight. Heard many in different places on our way
from Chesser Island to Starling Branch.
" 3 . Heard several in pond 5 mile west of Trader's Hill. Choruses in many

similar places. Associated with Bufo quercicus.
" II . Heard several at first and second crossings.
Aug. 8 . Along Chesser Island Road and Old Okefinokee Road heard quite a
few 8-1 1 P.M. Temperature 79°.
" II. Folkston-Chesser Island: Many immense choruses in wet grassy

places 8-12 P.M.
" 13. Commonly heard from St. Mary, Ga.-Chesser Island.

In 192 1 and 1922 the prevailing temperatures when we heard the species
in chorus frequently were from 'jo°-'jg°. An average of 50 temperatures taken
when the species was calling gives 74°. The most common range was from
7 1°-76°. The big choruses almost invariably came after a big rain. Then the
wet grassy places teemed with Httle Pseudacris ocularis. In 192 1 on May 20
we had an immense chorus, they were calling during the day. The tem-
peratures were ■jo°--j2° and the activity came after a rain. On June 24 and
25 after a rain they were common. Temperatures of 'jo°-'j6°, 72°, 73°, 76°
and much higher in mid-day prevailed. On July 15 a strong thunder storm
began. On this day and the following day Pseudacris ocularis without end
began. Temperatures from 73°, 76° and upward obtained. On July 29 we
were caught in a down pour on Black Jack Island, We were encamped in
the moist pine barrens. All about strongest chorus of Pseudacros ocidaris we
ever heard. Temperature about 73°.

In 1922 we had similar experiences. Humidity is first needed to send them
into breeding. The temperatures most prevalent are from 70° to 79°.

MATING

Coloration. In 192 1 "April 25 Dave Lee brought in some Chorophilus-\ike
forms." On April 28 we have these notes about these first specimens of 192 1
(our first since 191 2). "Three dead several hours. Maybe they are young
Hyla femoralis (they were not). Back from tip of snout brown with fine close
set series of grayish white spots giving creature a grayish back. Band from
tip of snout through eye over arm insertion half way or 2/3 way to groin.

Pseudacris ocularis 217

Band under eye to angle of eye pinkish bronze continued as broken series or
row of spots to arm insertion. Belly cream or yellowish with scattered black
dots widely spaced. Yellowish cream spots also scattered over belly and on
sides below black side stripe. Dark band on back of fore arm." Examination
of these prove them females. In spirits they are now (seven years later) with
the lateral stripe obscurely or slightly revealed.

"On August 18, 1922, at HilHard, Fla., (.7 mi. south beside Dixie Highway
found a little Pseudacris ocularis which is cinnamon or ochraceous tawny in
color." This proves an adult male with plaited throat.

Structural differences. In 19 12 we discussed this matter very briefly thus
"There are three males in the lot of ten, and each has indications of a promin-
ent gular pouch. These pouches are in a collapsed state and the wrinkles
are mainly in the lower throat region. All three have the breast fold promin-
ent; two are chestnut or bay and one olive or greenish. In one the throat is
very dark in color; in the others as light as the other ventral parts. There is
no decided enlargements of the fingers. Many of the males from 11.5 mm.
onward may have a plait parallel to lower jaw margin and somewhat mesad
of it. Then there is likely to be a collection of irregular folds in mid- ventral
throat instead of pectoral region. Sometimes it looks like a little triangle in
the mid-region of lower throat."

Some of the males from 11.5 mm. onward may have a plait parallel to the
lower jaw margin and somewhat mesad of it. There is sometimes one or more
often two mesal longitudinal plaits rarely three mesal plaits. Then there is
likely to be a collection of irregular folds in mid ventral throat ahead of pec-
toral region. Sometimes it looks like a little triangle in the mid-region of
lower throat. Sometimes two transverse plaits, rarely one, appear in al-
coholic males just ahead of pectoral fold.

Females have more tendency to have the lateral stripe prominent and the
dorsal band and intermediate dorsal obhque stripes to groin., i.e., 5 stripes.
Rarely a male has them but males more often have the lateral band absent.
In 1921 on June 23, we found a 14.5 mm. d^ ; on July 7, a 13 mm. cf , 14.5 mm.
cf, and 16 mm. 9 ; on July 8 a 13.2 mm. cf , 15 mm. cf ; on July 13-17, a 14
mm. cf , 14.8 mm. cf , 15 mm. cf, 17 mm. 9 ; on July 17 we took three pairs;
13.4 mm. cf , 14 mm. cf , 14 mm. cf ; 15.5 mm. 9 , 16 mm. 9 , 16 mm. 9 .

The measurements of one lot of 14 adult males are: three, 11.5 mm.; five
12 mm.; five 13 mm.; one 13.5 mm. The measurements of a group of four
ripe females are: 12 mm., 13 mm., 13.5 mm., 17.5 mm. Some of the freshest
material (U.S.N.M. 71034-41) are 8 frogs collected by Mr. Gerritt S. Miller Jr.
and C. R. Aschemeier, Apr. 3, 1926, at Gainesville, Florida. They are all
males with inflated throat sacs preserved in not entirely collapsed state. They
measure: two 14 mm., three 14.5 mm., two 15 mm., one 15.5 mm.

Thus we have externally apparent males from 11. 5-15. 5 mm., and females
1 2-1 7.5 mm. The males have two nodes 13 mm. and 14.5 mm., possibly ist-
year-olds and 2nd-year -old frogs. Surely this species begins breeding when
one year old if not sooner. The average of 35 adult males is 13 mm., also its
greater mode, the average of nine females is 15 mm.

2i8 Frogs of the Okefinokee Swamp

Duration, night or day. Pairs mated in laboratory the night of May 26,
192 1, were in embrace next day without ovulation. They were captured at
night and mated at night.

Amplexation {Normal, abnormal). The first amplexation came May 26,
1921. Several males and females were captured in the evening. "Put all
Pseudacris together. Soon two pairs were mated, one pair of male and
female. . . . Typical axillary embrace. They kept thus until we reached
camp and were flash-lighted without disturbing their amplexation."

On May 21, 192 1 "Caught a lot of Pseudacris males in the pond in the
afternoon. Two males amplexated. It was an axillary embrace." On May
26, 192 1 "put many Pseudacris together. Soon two pairs mated — One was a
female with two males embracing, one male holding the female axillary fashion
and other male holding the mating male axillary fashion. They kept thus
until camp was reached and were flash-lighted."

OVULATION

Habitat. The search for this life history was long and at times dis-
couraging. On May 26, 192 1 "we went for oak toads and Pseudacris. In
pond here the Pseudacris are gone from damp grassy sedgy places. Have they
reached the pond? Found only one in this area where for ten days they were
common." "Pseudacris often in little islands of grass mats where grass
extends 1-2 inches above water. Principally on little leaf islands with a black
gum or so and one or two overhanging bushes; on wet old leaves of former
years, also along edges of pond north and east side in grass or on leaves around
black gum trees. On one grassy log a male was calling; a foot or less away
was a female and not far from her another female. Apparently the females
are beginning to approach the water. Have seen a few females in the water.
Put all the males and females together. Soon two were mated." The follow-
ing morning. May 27, 1921, we noted: "No eggs as yet." On June 4 we went
to Chorophilus pond for habitat pictures. On mossy logs were Pseudacris.
Looked faithfully for a pair but did not find them."

On June 25, 192 1, "found a big congress. Are they finished breeding or
at height of breeding or what? Am afraid this form is so tiny it has slipped
by me."

Not until July 16, 192 1, did we find mated pairs in the field. From lab-
oratory mated pairs no eggs had come. "While searching for coatbets (Hyla
gratiosa) I found a pair of Pseudacris ocularis on a mossy log amongst Wood-
war dia fern. Tried for it and missed it. Later it leaped to a fern stem where
we caught it. In a stand of ferns next to a tree with hurrah bushes (Leucothoe
racemosa) were two pairs. Later Harper caught a pair but alas lost the male!
One or two more females were taken and several males. The pairs laid by
the next morning."

Period. On the basis of mating we have mated pairs from May 26 to
July 16 in 192 1 . On the basis of calling we have records from May 16 to 20 —
August 18 and some of the August records are of immense choruses in mid
August. In 191 2 a female taken June 16 was unspent. In 192 1 three females

Pseudaa'is ocularis 219

taken April 24 were spent; on May 23 three or four females were found un-
spent, and numerous ripe females and pairs were taken July 16. This implies
ovulation from April 2 4- July 16.

On the basis of tadpoles we have mature and transforming tadpoles by
June 2 8- July i, 1922, July 8 and July 19, just when we recorded heavy egg-
laying the year before. This imphes egg-laying in May or last of April.

In 1923 (1923, p. 34) we gave this species as one of five which started to
breed from "May 15 on to June ist." In our notes we variously have it "May
25- ," "May 26- July 18," "May i6-August 13" and in 1924 (1924,

p. 378) it appears May i6-August 21. This last period about represents
our knowledge of it in 1924 if we do not consider spent and unspent females
and transforming tadpoles. These latter send the ovulation at least to April
24 if not before. April 24-August 21 are the extremes of our evidence. This
species being a Pseudacris we might suspect it of appearing early but we dare
not prophesy breeding before March and possibly it may extend through
September to October i .

Temperature and humidity {See voice discussion). The only record of
positive ovulation came from July 15 and 16, 192 1 when temperatures from
73° and 76° obtained. This record comes after a hard rain. We have mating
records from 72° upwards. As determined under the topic of voice, ovulation
probably comes primarily after downpours when temperatures are usually
7o°-79°.

Egg-laying process. In locahties they choose even more shallow places
than Hyla crucifer, but were the author to show the eggs fastened to a stick I
doubt if few in America could tell them from Hyla crucifer eggs laid in cap-
tivity. We got the same captive complements as in this species. They lay
in somewhat similar places, have more or less similar individual eggs. We
will not be surprised to discover they lay their eggs in a similar way. In fact,
we suspect they belong nearer Hyla crucifer than near the other Pseudacris
species or Hylagratiosa, Hyla squirella, Hylaandersonii in egg-laying method.

EGGS

Attachment, dangers. One specimen, a female (No. 7 114), the specimen
where the lateral stripe breaks into mottlings, has large ripe eggs, which are
astonishingly large for the size of the frog. This specimen is only 15.5 mm.
long yet its eggs are .8-1.0 mm. in diameter. A slight examination of the egg
content shows 50 for either side is high and a complement of 100 eggs is the
lowest record for any North American Salientia. This female was taken on
Mixon's Hammock June 16, 19 12, in some overflow pools with spring frog and
toad tadpoles and with transforming cricket frogs. The middle of June is
much later than the breeding period for the larger northern Chorophilus
which are amongst the first to breed. These small males 12-13 mm. with
gular pouches apparent and this ripe female 15.5 mm. with ripe ova, very
strongly discredit any belief that these creatures are young forms of a larger
form. (Above note of 1912).

2 20 Frogs of the Okefinokee Swamp

In 192 1 these pairs of July 16-17 were captured in water from 3-12 inches
in depth. Their eggs hatch quickly and probably would not have been caught
by quick drying had they been laid naturally. Many eggs in small pools
might be caught except that ovulation usually comes at rainy periods.

The eggs laid the night of July 16-17, 192 1, were mainly attached to
sticks. "The three pairs were put into containers; two pairs in the photo-
graphic jar; the other pair in a round collecting bottle, all the equipment we
had available. Into each we put one or two sticks. The eggs are strewn
along on the sticks as in Hyla crucifer. Two masses of the three complements
were somewhat like normal Pseudacris masses (like P. triseriata-feriarmn-
nigrita group) of western New York and Texas forms. The eggs themselves
look like those of Hijla crucifer. The viteilus is brown above and creamish or
yellowish on lower pole. Ten of the eggs are floating. Quite a few single eggs
on the bottom of the jars. Must lay submerged eggs attached to grass, fern
and other herbaceous stems.

The next morning in this pond "in water 3-6 inches deep with dip net
discovered eggs which seemed Pseudacris ocularis eggs (like those in camp).
These eggs are harder than those of Acris.^^

Egg mass, egg description {See general account of eggs). The first eggs laid
in camp July 16-17, 192 1, were roughly measured. "The vitelli measured
from .6-.8 mm.; the outer envelope from 1.4-1.8 mm. With my lens I could
see no inner envelope. Egg reminds me of that of Hyla crucifer. Viteilus
brown and cream color."

In 1923 (1923a, p. 406) we gave their "eggs laid singly." In 1923 (1923b,
p. 34) we write '^Pseudacris ocularis emit single eggs on the bottoms of ponds
and on vegetation." In 1924 we (1924, p. 376) place the eggs in the category
"envelope single. Envelope 1.2 to 2.0 mm. Viteilus 0.6-0.8 mm. Egg com-
plement, 100. — Pseudacris ocularis."

Mrs. Wright's laboratory notes on 17-20 eggs are: Egg viteilus .8 mm. or
less; viteilus .6-8 mm.; viteilus mode .8 mm., average .75 mm.; viteilus brown
above, cream below; no inner envelope; envelope 1.2 -1.8 mm. in diameter;
envelope average 1.36, mode 1.4 mm.

HATCHING PERIOD

On the night of July 16 from 8-10 p.m. we found 3 or 4 mated pairs in a
pond on Chesser Island. Each pair laid eggs that evening. On July 18, we
have the note "The Chorophilus eggs laid night of July 16-17 hatching by
3 P.M. of the i8th. Fast work." This means a maximum of 42 hours if laid
soon after 10 p.m. or if not laid until 4 or 5 a.m. July 17, a maximum period
of 35 or 36 hours. These were in ordinary collecting bottles and petri dishes.
The temperatures for those days for surrounding stations were: minima 68°-
70°, average 72°; maxima 87°-92°, average 90°. The maxima probably were
very effective in the petri dishes.

MATURE TADPOLE

Color description from life {June 30, July 8, 1922). Dorsal color citrine
drab or deep oUve. Over the dorsum of the body are definitely scattered

Pseudacris ocularis 221

distinct black spots. Lower belly a block of orient pink or orange pink. Some-
times this is broken into a mottled arrangement when the gill region and sides
are reached. Area around the mouth and an extension backward on each
side martius yellow. Heavily speckled on under side of developing hind legs
or femur when two legged stage is well started.

Tail. Lower edge of muscular part for the basal one-half a line of martius
yellow. The upper edge a rim of muscular part apricot buff to apricot orange
or rufous to ferruginous. This color extends on to the top of the body and
gives it the same color dorsally as upper musculature. The middle of the
muscular part is with a prominent sharply defined band of chestnut brown
or black at the basal half or two-fifths of the tail and hazel or cinnamon
rufous onward to tip. This dark middle stripe extends onto the body in
transforming individuals, becomes the dark stripe either side of the median
dorsal stripe. From mature tadpole to transformation the three-striped
(light, dark light) arrangement obtains, the lower stripe being least con-
spicuous. The rim of the tail crests is with large blotches. The interval be-
tween the crest rim and the musculature is translucent and without spots or
dots.

Iris black, citrine drab or deep olive finely speckled with empire yellow or
bittersweet orange.

In alcohol the broad brown tail band follows on to body to back of the
eye. Above brown band is a clear white edge which extends on to the body
and around the edge and then forward. Below brown band is a light band.

General appearance. Tadpole quite small (23 mm.) full and deep bodied
(in general appearance somewhat like H. crucifer). Tail quite long, tail tip
sharply acute or even acuminate. The dorsal and ventral crests about equal.
The dorsal crest not deeper than tail musculature and extending on to the
body to the vertical halfway between eye and the spiracle. Spiracles sinistral,
directed backwards and upwards, far below, lateral axis, the spiracular open-
ing prominent elliptical round. Eye on the lateral axis, dorsal aspect on the
outline and therefore visible in the ventral aspect as well. Anus dextral, open-
ing on a level with the lower edge of the ventral crest. Muciferous crypts not
distinct.

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
labial fringe for about 2/9-1/4 of the length of the upper fringe. The horny
beak is contained in upper fringe 2.0 times in the upper fringe. The median
space between the second lateral upper labial rows of teeth long, 3-4 times
the length of either lateral row. Inner papillae sparse or absent on upper
labium and opposite the anus or horny beak. At ends of the second lower
labial teeth row two or three rows. Sometimes on sides two rows of labial
papillae from upper fringe down. The papillae do not extend under second
row to the end of the third row of labial teeth as in H. crucifer. The second row
of papillae across the lower labial border is about equal to the second row of
labial teeth. The third labial row of teeth is short, 1/4-1/3 the length of the

2 2 2 Fi'ogs of the Okefinokee Swamp

first or second row of lower labial teeth. The first and second rows about equal
and about two times the horny beak in length, upper fringe somewhat an-
gulate in the middle.

Measurements. Length of body (7.8-9.0 mm.) in tail (12-14. 4 mm.) 1.33
-2.0, average 1.58. Width (4.0-4.8 mm.) of body in its own length 1.3-2. i,
average 1.65. Depth (3.8-5.0 mm.) of body i. 0-1.4 in body width, average
I.I 7. Depth of body 1.72-2.2 in length of body, average 1.90. Depth (3.2-
4.0 mm.) of tail in length of tail 3.1-4.4, average 3.58. Muscular part (2.0-
2.2 mm.) 1.7-2.2 in depth of tail, average 1.88. Spiracle 1.7-2. i nearer base
of hind legs or vent region (2.8-3.8 mm.) then the tip of the snout (4.8-6.8
mm.), average 1.675. Spiracle to eye (3.0-3.4 mm.) equals spiracle to base of
hind legs or vent. Eye to tip of snout (2.6-3.8 mm.) about equal to eye to
spiracle. Nostril 1.25-2.0 nearer eye (i. 2-1. 6 mm.) than snout (2.1-2.4 mm.),
average 1.5. Mouth (2.0-2.8 mm.) usually equals internasal space (2.0-2.8
mm.) Mouth contained 1.5-1.9 (average 1.66) in interorbital distance (3.0
-4.2 mm.). Internasal space contained in interorbital space i. 3-2.1, average
1.63.

The dimensions of the largest tadpole are :

Total length

23.0

Spiracle to vent

3-8

Body length

8.8

Spiracle to eye

3-0

Body depth

5-0

Eye to snout

3-0

Body width

5-2

Eye to nostril

1.2

Tail length

14.2

Nostril to snout

2.2

Tail depth

3-2

Mouth

2.2

Musculature of tail

2.0

Interorbital distance

3-4

Spiracle to snout

6.8

Internasal distance

2.2

General remarks. In 1923 (1923, p. 406) we characterized the tadpole
thus "Eggs laid singly. . . . Pseudacris ocularis scattered distinct black dots on
body, black band on musculature of tail, clear intervals between musculature
and rims of the crest." In 1928 we summarized it thus "Musculature with a
distinct brown lateral hand with light area below; crests usually clear with fine
scattered fleckings, sometimes with fleckings gathered near outer rim; one or
two rows of papillae below the hind lower labial row of teeth ; nostril to eye in
nostril to snout 1.25 -2.0; depth of tail in tail length 2.9-4.4, average 3.4,
3.55; spiracle 1.15-2.1 nearer vent than snout, average 1.5-1.875; median
space between second upper labial row 2 1/2 or 3-7 in either lateral portion;
horny beak in upper fringe, 1 5.-2.0. (Characters above shared with Pseud-
acris of Raleigh). The characters given for Pseudacris ocularis are: ''Dorsal
crest to vertical midway between spiracle and eye; spiracle equi-distant from
eye and vent; spiracle 1.7-2. i, average (1.875) near vent than snout, mouth
and internasal space equal; one row of papillae below and third lower row of
teeth; third lower labial row of teeth .33 of the first lower row; dorsum of

Pseudacris ocularis

223

body in life with definite scattered black spots ; musculature with three bands,
apricot buff (light), chestnut brown (dark) martius yellow (light); first and
second lower labial rows 2.0 greater than the horny beak; horny beak in upper
fringe, 2.0; median space between second upper labial row of upper labial
teeth 3-4 in either lateral portion. Eggs single."

LARVAL PERIOD
We never raised this species from egg to transformation. Too many in-
vestigations were going at once to get them past the adult tadpole stage and
such laboratory tadpoles reared for identification could give us little real idea
of the larval period. The earliest we know the species to have bred is April 24;
the earliest we have taken tadpoles transforming is June 30. I believe the
difference between those two dates or 67 days is an average period for the
species. The first stray choruses, or mating we have observed came in the
latter half of May and the bulk of our transformation records is the first of
July. This presumptive evidence would give about 45 to 60 days for larval
period. Forty-five to seventy days may represent the larval period. More
evidence is needed.

TRANSFORMATION

Period. On June 30, 1922, in Saddle-bag Pond (near Chesser Island School
U. S. Geol. Moniac Quad.) "found these tadpoles transforming in Wood-
wardias (ferns) around the edge of this cypress pond. Also in open pond,
slightly so. Transformed ones have median stripe most conspicuous, the
stripe of the tail continues on the body as a stripe either side of the median
stripe and the stripe through the eye least conspicuous. The ground color of
the adults is more or less prominent in the dorsal of the tadpoles. There is
a prominent black band on the muscular part of tail. Very strongly defined
black band, this is with light band below and a less conspicuous one above
for 3/16-1/8 of an inch. This last extends on to the body." From this same
pond tadpoles taken July 6 transformed July 8.

On July 18, 1922, "around the west edge of Murray Bay in the more or
less flooded area (at times not now) of gums and pines found many transformed
Pseudacris ocularis. They were difficult to catch because of minute size,
fear of crushing them and their marvellous abihty to jump. In grassy edges
these transformed frogs and adults are also present."

On August 18, 1922, "at HiUiard, Florida, beside the Dixie Highway found
Httle Pseudactis ocularis which is cinnamon or ochraceous tawny in color."

Size. On July 2, 1922, we took six transforming and transformed frogs
of the following lengths and characters: Two, 9 mm. transformed; one, 9 mm.
and only 2 mm. stub; one 11 mm. and 2.5 mm. stub; one, 9 mm. plus 1.05
mm. tail, frog-like appearance; one 9 mm. plus 1.25 mm. tail, tadpole ap-
pearance,— 9-1 1 mm. in transformation size. Twenty to twenty-five tad-
poles taken at the same place and many with prominent hind legs have body
10 mm. I beheve the 11 mm. transforming specimen abnormally large for
this species.

224 Frogs of the Okefinokee Swamp

Two taken July 8, 1922, were: one fully transformed 8 mm.; one 8.5 mm.
(with tail 1.35 mm.). On July ig in another place we found five transformed
and transforming individuals and 19 mature tadpoles. The five were: three,
8.5 mm.; two, 9 mm.

On July 18, 1922, in Murray Bay found 29 transformed Pseudacris ocu-
laris; two, 7 mm.; seven, 7.5 mm.; nine, 8 mm.; six, 8.5 mm.; five 9 mm.

All in all the size seems to range from 7-9 mm., one specimen 11 mm.
probably being abnormally large. Of 41 specimens two are 7 mm.; seven
7.5 mm.; ten 8 mm.; ten 8.5 mm. and twelve 9 mm.; 8 to 9 mm. being the
average and modal size.

GROWTH

At transformation these pygmies are 7-9 mm. in length. On July 18,
1922, we took 29 transformed frogs from 7-9 mm. and five adults: two males
1 1.5 mm.; one male 12 mm.; one male 13 mm.; one female 13 mm. — two size
forms 7-9 mm. and 11. 5-13 mm., the latter doubtless one year old. We have
some isolated specimens of males 13.5 mm . I n 1 9 1 2 the males were 12-13 ni™ .
one female 15.5 mm.

Cope gives .62 of an inch for total length. Our female 17.5 mm. is .70 of
an inch and may represent an approximation of the maximum, though I
am positive I have seen larger specimens.

In 192 1 on May 16, four 12 mm., four 13 mm., and one 11.5 mm. males and
three females, 12 mm., 13.5 mm., and 17.5 mm. were taken. On June 23,
one male 14.5 mm. was taken. On July 7, we have 13 mm. cf, 14 mm.cf ,
16 mm. cf. On July 8, 13.2 mm.d^ and 15 mm.d^. On July 13-17 we took
i4mm.<:f , 14.8 mm. cf, 15 mm.cf , 17 mm. 9. On July 17, one more set of 3
pairs give 13.4, 14, 15 mm. cfs and 15.5, 16, 16 mm. 9 s. Miller's group
(U.S.N.M. 7(034-41) of eight males 14- 15.5 mm. look comparable to the
17.5 mm. female above and our 1912 female of 15.5 mm. or the 1921 i4mm.-i7
mm. material.

The evidence seems to point to 7-9 mm. for transformation; 11. 5-13. 5 for
I st-y ear-olds; 14-17. 5 mm. for second-year-olds. Possibly 11. 5-17. 5 mm. all
represent first-year-olds.

FOOD

We have made no study of this phase nor has any other naturalist. The
food of these brownies of our frog species would be very interesting to study;
must be minute insects, spiders, etc. Those who wish to work with the
smaller insect forms ought to find this species a good collector for entomolo-
gists.

ENEMIES

On May 17, 192 1 on Billy's Island we found a fine Thamnophis sackenii
(Ribbon Snake) where little chorus frogs were in grassy, sedgy, pipewort,
heath cover. This species must be a common prey for many snakes of the
ponds, bays and moist pine barrens and for other animals as well.

Pseudacris ocularis 225

AUTUMNAL APPEARANCE

We know nothing of the activities of this species beyond the first of
September. Doubtless it is active most of the year except for mid-winter.

AFFINITIES

Holbrook remarks that Hylodes ocularis "is closely allied to Hylodes
gryllus in its habits, choosing the same damp places for its abode, — and like
Hylodes gryllus, too, it has the power of adhering to smooth surfaces, though
not so perfectly so as the Hylae.'" It is quite arboreal at times on boles of
trees, etc., far more arboreal than Acris.

Holbrook (1843, p. 38) remarks: "I have chosen the specific name ocularis
for this animal, from the black spot along the eye; and this has been done the
more willingly, as it is possible this animal may in the end prove to be the
Hijla ocularis of Bosc and Daudin, which LeConte thinks, however, is most
probably only a variety of the Hyla squirella; and yet, as this animal is not
infrequently found on low bushes, or leaps upon them when pursued, it is not
possible that Bosc might have supposed it to be a real Hyla. Should it then
turn out to be the Hyla ocularis of Bosc, an additional synonym will be saved;
but if it is an undescribed animal, which I beheve, then the demonination
Hylodes ocularis is well enough, as it belongs to another genus."

LeConte (1855, p. 429) places it in Hyla ocularis following Hyla pickeringii
in his account. He credits it to Daudin. "From the small size of this and the
preceding species, {Hyla pickeringii) the web between the third and fourth
toes is not very perceptible." LeConte in this article recognizes ocularis as of
Hyla, ornata and nigrita as of Chorophilus.. A year before he recognized
Chorophilus with nigritus, Helocaetes with feriarum, triseriata and clarkii.

In some ways were it not for the tadpole coloration, the single eggs, self
color and some structural characters one might be led to consider P. ocularis
apart from the other Pseudacris which are of larger size, more spotted backs
and with egg masses. Pseudacris septentrionalis, P. triseriata, P. nigrita, P.
feriarum generally so far as known and from my own experience tend toward
the laying of egg masses rather than single eggs. Our life history material on
Pseudacris ornata is too fragmentary for positive assertion on this point.

Miss M. C. Dickerson (1906, p. 157) writes: "The Chorophilus material is
confusing and insufficient to settle any problems. . . . However, the whole
subject is in need of investigation and is open to revision." In 1923, Noble, her
successor, wrote on the Generic and Genetic Relations of Pseudacris, The
Swamp Cricket Frogs "basing his conclusions largely on a detailed considera-
tion of Pseudacris ocularis. His conclusions are that "The species of Pseud-
acris must be referred to Hyla, although all species are distinguishable from
American Hylas on external characters."

In egg characters it seems to fall nearer Hyla crucifer in size of eggs,
envelopes and vitelli where LeConte apparently placed it in Hyla. In other
ways it is not far removed from Acris gryllus with which Holbrook placed it
as of Hylodes. Possibly Acris lays masses at times (single usually) like other
Pseudacris species.

2 26 Frogs of the Okefinokee Swamp

In tadpole coloration Pseudacris ocularis has the brown lateral band of the
other Pseudacris species. In tadpole pattern it is nearer Hyla crucifer than
Acris gryllus. In labial teeth it has a formula 2/3 not 2/2 as in Acris gryllus
regularly and sometimes in Hyla crucifer. The latter however is also 2/3 and
some Pseudacris tadpoles (other species) may have 2/2.

In adult coloration it approaches more nearly in some individuals (five
stripes) the Pseudacris type rather than that of Hyla crucifer or Acris gryllus
but unlike most individuals of Pseudacris species it is usually self color on the
back. The author has seen in the field many fine Pseudacris from New York
to Brownsville, Texas, and this species in many ways seems the most distinct
species of the group unless it be Pseudacris ornata. Each has been poorly
understood. Pseudacris ocularis proves a common form wherever found.
So, too, Pseudacris ornata doubtless will prove not so rare. Had the average
person lived and played with this species from 1912-1922, as some of us have,
he might conclude Pseudacris ocularis the most distinct and possible aberrant
■species of the group both in diminutive size, in self color (others sometimes
are though not always so), in singly-laid eggs and other impressions one gets
from the species. With only Pseudacris ocularis and Pseudacris ornata at
hand one would question whether to put them together if he had not been
schooled in the history of the group. Furthermore, though the present author
has worked out the life history of three or four species of Pseudacris in twenty
years he dare positively assert that "The species of Pseudacris must be referred
to Hyla. . . ." In the same paper we have "Future work may show that P.
septentrionalis, P. triseriata and P. feriarum are closely allied and possess
characters in common, distinguishing them from the more Hyla-like species
of the genus." Stejneger and Barbour in adopting their course in retaining
Pseudacris until this central core of Pseudacris are carefully worked out — life
histories, field impressions, anatomy and all — take the more conservative
course. All the collections in the U. S. A. will not clear it up like actual field
experience with each form. I would not be surprised, however, if some, if not
all of Pseudacris might be eventually referred to Hyla as Noble suggests, but
why hurriedly consign them all on the detailed study of one possible outlying
species. Nevertheless Pseudacris ocularis is as near Hyla crucifer as any hylid.
It is Sabalian in distribution and Hyla crucifer more northern in general. Or
it may be a good Pseudacris coincidental within the range of other Pseudacris.
Apparently Pseiidacris is largely an upper Austral group (except possibly for
P. septentrionalis).

BIBLIOGRAPHY

1882 Boiilenger, G. A. Catalogue of Batrachia Salientia S. Ecaudata in . . . B. M. 2nd

edit. London 1882, pp. 333, 334.

1926 Brimlev, C. S. .Journal Elisha Mitchell Soc, Vol. 41, Nos. 3 & 4, Apr. 26, p. 82.

1875 Cope, E. D. Check List— List Batr. & Rept. N. Am., 1875, p. 30.

1Q13 Deckert, R. F. Copeia, Feb. 14, 1914, No. 3, p. 3.

1915 . Copeia, July 27, 1915, No. 20, p. 23.

1921 . Copeia, Mar. 15, 1921, No. 92. p. 22.

1922 . Copeia Nov. 20, 1922, No. 112, p. 88.

1906 Dickerson, M. C. The Frog Book p. 162.

1838-42. Holbrook, J. E. North American Herpetology, ist ed. II, 79, pi. XIV, 1838;
2nd ed. IV, pi. 33, 1842.

Pseudacris ornata

227

. Cat. Reptiles of Georgia. In White, Geo. Statistics of Georgia, Savan-
nah, 1849. Appendix p. 15.

1855 LeConte, John Proc. Acad. Nat. Sci. Phila., Dec. 1855, p. 429.

1924 Myers, G. S. Copeia, June 30, 1924, No. 131, pp. 59, 60.

1923 Noble, G. K. Am. Mus. Novitates Apr. 23, 1923, No. 70.

1923 Stejneger, L. & T. Barbour. A Check List of North American Amphibians and
Reptiles. 2nd Edition, Cambridge, 1923, p. 28.

1923 Wright, A. H. Copeia, Feb. i, 1923, No. 115, p. 34.

1923 Wright, A. H. and A. A. Wright. The Anatomical Record Jan 30, 1923, Vol. 24,
No. 6, p. 406.

1924 Wright, A. H. & A. A. Wright. The American Naturalist, July-August, 1924.
Vol. LVIII, p. 379.

Pseudacris ornata (Holbrook)

(PI. I Fig. 8)

ORNATE CHORUS FROG. ORNATE TREE FROG.

Holbrook in 1842 credits this form to South Carolina and DeKay followed
him in giving South Carolina as its range. In 1849 Holbrook gives Georgia
as also a part of its range. LeConte 1855 (doubtless on Riceborough evi-
dence) says it ''inhabits Georgia, very common in the pine barren ponds."

Gunther (1858, p. 29) gives Cystignathus ornatus with "Body and head
short as in Hyla viridis; skin smooth; tympanum distinct; tongue nearly cir-
cular; vomerine teeth in two rounded groups behind the inner nostrils. Dove-
colour above with oblong spots of dark brown, margined with yellow.

a. Adult. Georgia. From Mr. Cuming's Collection. Differs in having the
back uniform olive-colour; sides with three black spots, viz., a streak from the
eye to the shoulder, an oval spot in the middle of the sides and one or two
smaller ones on the loin."

(See discussion under P. occidenialis). We did not take Pseudacris ornata
in the swamp in May-June 1912, nor in April-Aug. 192 1 nor in June-August
1922. We have seen it at Helotes and Beeville, Texas in 1925. C. S. Brimley
(see P. occidenialis) received it from Green Cove Springs, (1898) and Hastings
(1901) Florida. In 1930 and 1931 we have received it in February or early
spring from O. C. Van Hyning of Gainesville, Florida.

Hyla andersonii Baird

(PI. II, Fig. 3; X Fig. i; XII, Fig. 3; XV Fig. i; XVII; XXV)

COMMON NAMES

Anderson Tree Frog. Anderson's Tree Frog. Anderson's Hyla. Ander-
son Tree Toad. Green and Yellow Tree-toad.

(This species and Rana areolata are two species the author has studied the
least. Noble and Noble, 1923, have treated most of the essential features of
this species. It is included here to make comparisons, and to add our im-
pressions of it at Lakehurst and North CaroHna and as a captive in 1906- 1908.
It is also a possible Okefinokee anuran.)

2 28 Frogs of the Okefinokee Swamp

RANGE

Check list. "Type Locality: Anderson, South Carolina.

Range. White cedar swamps in Southern New Jersey". Stejneger and
Barbour, 1923, p. 29.

Supplementary records. The Check List (above) appeared about coincident
with the pubhcation of the extended studies of Noble and Noble (1923). In
view of their detailed observations the present writer will not attenpt a long
account of this species except to summarize some topics so that this species (a
possible hypothetical species of the Okefinokee Swamp region) may have the
same form treatment as the others.

The writer, unlike some of the Philadelphia or American Museum groups,
has not had the continued experience with this species some of these gentlemen
have. Nevertheless, he has seen and recorded this species in North Carolina
and New Jersey near the two extremities of its range. After Baird's descrip-
tion (1854, pp. 60, 61) all the records from 1854-1904 are from the Philadelphia
(and its environs) group of naturalists (Leidy, Cope, Peters, Abbott, Moore,
Stone, Fowler, Rehn, Klein, etc.). From 1898 onward the New York group
(W. T. Davis, James Chapin, W. D. Miller, C. L. Camp, K. P. Schmidt, G. K.
Noble, G. S. Myers and R. Sackett), led by W. T. Davis have followed the
trail until its habits are well enough known for life-history studies. Early in
that search James Chapin was associated with W. T. Davis and our first ac-
quaintance of this species alive came through two specimens "sent July 21,
1906, to Dr. Burt G. Wilder from James Chapin, these he collected at Lake-
hurst, N. J." We mention this because Fuertes (who recently died) painted
one of these frogs — one of his only amphibian paintings (now in possession of
Professor H. D. Reed). Besides from 1906 specimens of these gentlemen,
came Miss Dickerson's excellent plate of a female (Dickerson 1906, Plate VII).

It is interesting that the first two, W. T. Davis and James Chapin (who
first began 1 904-1 906 studying this form) should be the first to record it from
North Carolina (1920) and we (1922) who received first five specimens (1906)
from them, should independently (1922) record this species in North Carolina
35-50 miles southwest of Southern Pines, their locahty of 1920. Our record is
midway between Rockingham, N. C. and Cheraw, S. C. on the North Carolina
side of the State Line at Everett's Pond. Southern Pines country is partially
of the same river drainage (Pedee River). If any one wishes to record it in
South Carolina he can follow the Pedee down from Everett's Pond and no
doubt easily find it.

The record from Anderson, S. C, has always bothered the writer. In 191 7
we went through Anderson and had no opportunity to collect at this place.
Anderson is twice as far from the coast as Southern Pines, N. C. or Cheraw,
S. C. is from the coast. Possibly the species extends that far inland along the
Savannah River. No fife zone map can show all the river valley extensions
inland. Anderson's tree frog is a lower Austral (Austroriparian, Upper Coastal,
etc.) form, and Anderson, S. C. is in Upper Austral (CaroHnian, Piedmont,
etc.) country. If such terminology be employed it may be well to remember

Hyla andersonii 229

that C. S. Brimley and F. Sherman, Jr. (1908) send the lower Austral Hne from
Raleigh to Charlotte, N. C. or farther west. In this event possibly Anderson's
tree frog is not from Anderson.

Of its northernmost range, W. DeW. Miller (1906, p. 68) writes: "About
eight years ago James Chapin and I found this beautiful Tree Frog at the
Runyon Pond two miles south of Sayreville, Middlesex County. Since that
time I have found it at several other localities in the same region — one mile
south of Old Bridge, about one mile southeast of Browntown and at Freneau
near Mattewan. These localities are all in the sandy pine barren 'island'
north of the Pine Barrens proper. Thus this species extends northward to
within three miles or less of the lower Raritan River, its range coinciding at
this point with that of the Carolina Chickadee."

Of its range northward, W. T. Davis, two years later (1908, p. 49) writes,
"In addition to the notes on this species printed in the American Naturalist,
it may be added that it occurs still further north than Lakehurst and Farming-
dale, and has been heard by the writer at Jamesburg on the South River.
Messrs. Miller, Chapin, and Skinner have found the frog at Runyon, also on
the South River. It no doubt occurs as far north as the Raritan."

If we call it a pine barren's form we must remember it is not recorded on
Long Island nor westward of Anderson, S. C, in Georgia, Alabama, etc. If it
be termed a sphagnaceous tree frog why is it not in the Dismal Swamp, Great
Alligator Dismal Swamp (N. C.) or Okefinokee Swamp? On the basis of as-
sociated species Scaphiopus holhrookii extends into New England and south-
ward to Florida and Texas. Acris gryllus is not comparable in the southern
states and if not found in pine barren bogs in New Jersey, it is abundant in
such places and many other localities in the South. Even yet in spite of the
fact that Hyla andersonii outstrips it northward (See W. D. Miller), Rana
virgatipes seems its nearest associated species, though Rana virgatipes outstrips
it southward to Okefinokee Swamp.

In northward range Rana sphenocephala if recognized as distinct doubtless
about coincides in its northern extension with H. andersonii. Though not yet
recorded from New Jersey (1925) some meadow frogs from Lakehurst south-
ward look much like Rana sphenocephala.

After my experiences with Rana virgatipes in New Jersey, North Carolina
and Georgia the writer instinctively expects and looks for H. andersonii
wherever he hears Ra7ia virgatipes. True in Dismal Swamp, Craven Lakes,
Wilmington and in Okefinokee Swamp they are not yet recorded, but they
will doubtless be recorded near these places with intensive search. To know
the notes of frogs gives the best clue to possible new records for "rare" frogs.

GENERAL APPEARANCE

Cope (1887, p. 365) held it "in proportions and general appearance similar
to the Hyla arborea of Europe." In its stout body it differs from the more
slender Hylas, H. cinerea and H. cinerea evittata. By its lateral band, it has
its dorsal color more sharply defined from venter than in any other species of
our Hylids. Brimley (1907, p. 158) gave the "plum colored line along the
sides of body with yellow spots below it" as its distinct mark.

230 Frogs of the Okefinokee Swamp

COLORATION OF SPIRIT SPECIMENS

Fowler (1908, p. iii) in describing Viereck's specimen writes "After hav-
ing been in alcohol several years it has changed to a plumbeous or slate-gray
color above with paler or whitish borders to brownish of sides and dark
lateral stripes. The limbs through plumbeous above are only so to carpus
and tarsus, which are a pale brownish like entire under surfaces of limbs.
Throat, breast and belly pale or whitish." Leidy's specimen in alcohol faded
to "dull brownish above and whitish below." (Fowler, p. no). The author
quotes at above length because a Hijla eviitata or Hyla squirella might be
confused with an old specimen of Hyla andersonii. In one collection a group
of Hyla squirella was confused with Hyla andersonii, but the light upper lip
of Hyla squirella finally separated them. So also alcohohc specimens without
vitta and Une revealed are hard to separate from Hyla cinerea eviitata. One
specimen twenty years in alcohol has a light line hke Hyla cinerea but not a
straight line as in Hyla cinerea. Usually some indication of the vitta before
.and behind eye is revealed. One specimen preserved in 1906 is vinaceous
drab or purple, drab or livid brown on upper parts and throat region. Another
of same year is greenish glaucous blue or Niagara green.

STRUCTURAL CHARACTERS
This phase has had much attention and needs no repetition here.

MEASUREMENTS

(Recent Material)

We have no 20 and 28 mm. specimens. Our measurements of one male
36 mm., and two females 36 and 44 mm. respectively give Uttle idea of the
range of relative measurements.

Head to angle of mouth 1.16 (36 mm. cf) -1.2 (44 mm. 9) in width of
head; head to rear of tympanum 1.07 (36 mm. cf ) -1.2 (44 mm. 9 ) in width
of head; head to angle of mouth 3.0-3.5 in length of body; head to rear of
tympanum 2.77-3.3 in length of body; snout .92-.94 in first finger; eye .92-
i.o in first toe; eye 144-1.33 in snout; eye .8-.66 in tympanum; eye i. 3-1. 44
in first finger; tympanum 3.3-5.2 in intertympanic width; tympanum 18.6-
2.6 in snout; internasal width 1.0-1.125 in upper eyelid width; interorbital
width .80-.88 in upper eyelid width; interorbital width .80-.9 in internasal
width; interorbital width 2.4-2.9 in intertympanic width.

Forelimh. Forelimb 1.8-2.0 in length of body; forelimb 2.7-2.0 n hind
limb; first finger 1.5-2.76 in third finger; second finger 1.3- 1.2 5 in third finger;
second finger .85-7.5 in first finger; third finger .7 7-. 85 in second toe; fourth
finger 1.1-1.25 in third finger; fourth finger .7 5-. 7 5 in first finger; internasal
width 1. 5- 1. 5 in first finger; internasal width 1.75-2.0 in second finger; inter-
nasal width 2.25-2.5 in third finger; internasal width 2.0-1. 41 in fourth finger.

Hindlimh. Length i. 5-1. 41 in hindlimb; tibia 2.0 (44 mm.) in length;
tibia 3.0 in hindhmb; tibia i.i in forelimb; tibia .90 in hind foot; first toe
1.16-1.7 in second toe; first toe 1.66-2.6 in third toe; first toe 2.16-3.5 in

Hyla andersonii 231

fourth toe; first toe 1.8-2.7 in fifth toe; second toe 1.43- 1.5 2 in third toe;
second toe 1.85-2.06 in fourth toe; second toe i. 56-1. 6 in fifth toe; third toe
1. 3-1.34 in fourth toe; third toe 1.1-1.04 in fifth toe; fourth toe 1.27-.97 in
hind foot without tarsus; fourth toe i. 38-1. 2 5 in tibia; internasal width
1. 5-1. 2 5 in first toe; internasal width 1.7 5-2. 12 in second toe; internasal
width 2.5-3.25 in third toe; internasal width 3.25-4.4 in fourth toe; inter-
nasal width 2.75-3.4 in fifth toe.

HABITAT

(See Voice discussion for habitat notes)

In 1908 Mr. Davis (pp. 49, 50) presents some good habitat notes which we
quote in toto. "On the evening of July 19, 1907, I went hunting Hyla ander-
sonii at Lakehurst. The first one that I heard was in a small red maple and
about eight feet from the ground. This was quite high up, for they usually
climb only five or six feet from the ground. I could not reach the little frog,
nor could I bend the sapling that hung over a marshy place. I therefore shook
it suddenly, which caused the frog to jump into a still smaller maple. This I
could bend over, and it sat looking at me until I captured it with my hand.

''Of quite a different disposition was the next one I found. It also was in a
small maple, but when it saw me and I tried to capture it with my two insect
nets, it immediately jumped and as it was getting quite dark by that time, I
was unable to locate it on the ground among the undergrowth. A little later
I heard it singing again, and so went back to find that it had climbed up a
huckleberry bush that was covered with a considerable growth of Smilax
glauca. The leaves of the vine made it quite difficult to locate the frog, which
must have been near, for when I drew close it hopped on to a lower part of the
bush, and though I tried to capture it with my two nets, I was unsuccessful.
This frog was a most persistent singer, and so in a short time I went back to
the same bush up which it had climbed after being so rudely dislodged. After
much looking and with the aid of the moon, I located the frog on the flat side
of a Smilax leaf and brought the two nets together with the result that I
found the frog in one of them.

"The next evening I found one in a cedar tree in a swamp. When I got
around the tree through the bushes to where the frog had been, I found that
it had gone to the side of the tree that I had just left. As I approached
slowly it would jump from tree to tree, and from limb to limb, every now and
then stopping to sing when it heard the calls of its rivals, two of which were
in the same small swamp. In this way I slowly pursued the frog, being much
interested in its well-founded fear and its inability to keep quiet. It never
cHmbed over sixfeet from the ground so that I easily captured it when I desired."

In 1928 Dr. & Mrs. A. B. Klots located ten individuals and their notes
on position are apropos.

"The exact positions of ten individuals were located, of which seven were
captured. High-bush Blueberry tangles festooned with Green Briars made
further investigations in this line impossible. The individuals are here re-
ferred to by number.

232 Frogs of the Okefinokee Swamp

"Numbers i, 2, 3, 4 and 5 were in one group, within 50 feet of the main
'pike' from May's Landing to Hammonton. The ground was covered quite
evenly with Blueberry bushes from a foot to 18 inches high. Scattered Pitch
Pines up to 12 inches in diameter stood from 10 to 30 feet apart. The ground
was at most damp, and the only water nearby was a shallow pool about 30
feet away which probably dries up in the summer. Near the bases of the pines
stood taller Blueberry bushes, up to three feet in height. No. i was sitting
on the main stem of a small Blueberry bush, 18 inches from the ground and
six inches from the tip of the bush. A Pine stood i^ feet away. No. 2 was on
the leaf of a Blueberry bush, 2^ feet from the ground and i| feet from a Pine
trunk. No. 3 was on a little twiglet growing out from the trunk of a pine
3I feet from the ground. No. 4 was on the ground at the base of a Pine.
No. 5 was one foot from the ground, where the twig of a Blueberry bush lay
against the trunk of a Pine. All of the specimens in this group showed a
strong preference for the near vicinity of a Pine.

"Nos. 6, 7 and 8 were in a thicket of small Red Maples and high Blue-
berry bushes in a creek 'bottom'. No. 6 was on the main stem of a Blueberry
bush about four feet from the ground. No. 7 was similarly located. No. 8
was about six feet from the ground in a small Red Maple.

"Nos. 9 and 10 were in a thick tangle of high Blueberry bushes and
Sndlax. Both were near the tops of Blueberry bushes at least nine feet from
the ground. For fifty feet around none of the vegetation was any lower, so
it seems that these individuals climbed higher than is usual for the species
in order to be out in the open.

"Not all of the individuals were as tame as is generally noted for andersonii.
A number of individuals would not continue singing when the observer
turned the light on them or approached nearer than fifteen or twenty feet,
and so could not be located. A silent andersonii in a thick tangle of Blue-
berry bushes could give points on hiding to a very small needle in a very
large haystack. No females were taken."

FIRST APPEARANCE

The records of earliest spring appearance are the following: "a specimen
of Hyla andersonii obtained at Clementon, N. J., May 12, 1901, by Mr.
Henry L. Viereck"; (Stone, 1901, p. 342). Several down to Noble and Noble
(May 14) have recorded them in May. The reason adduced by Stone for its
scarcity also explains why earlier records in spring and later records in fall
are not available. Stone held (before Davis proved it abundant) that "though
probably of restricted distribution the species would no doubt prove more
abundant if specially sought for, the comparative remoteness of the New
Jersey barrens, where most of the specimens are found, and the retiring habits
of the animal both tending to make its detection difficult". (Stone, 1901, p.
342). Of course, this species appears earlier than May 12,

Hyla andersonii 233

GENERAL HABITS

Metachrosis. This species is not recorded as capable of a wide range of
color variation. Miss Dickerson (1906, p. 133) records that her lone female
specimen might pale or deepen in its green or take on a tinge of brown. Noble
andNoble(i923,pp.432,433)onthebasisof their experience with four or more
pairs and others seen or taken noticed that light, excitement or humidity
might change their color. The first and last factors are most analyzable and
their results accord in the main with most species. The writer suspects that
time will reveal as marked variation in general color in this species as in
Hyla cinerea. Hyla andersonii seems truly more constant than Hyla squirella
but once upon a time green Hyla squirella were pronounced unknown. Out-
right statements concerning color are dangerous. For example two males of
Hyla andersonii were received July 21, 1906. Each had a similar green
backed livery. One we fixed at once in alcohol. Today it is a shade of dark
green. The other was kept until November 27, 1908, when it was fixed in 5%
formol. It is now in alcohol and is purplish drab.

Variations in color. (See Hyla cinerea).

General habits. Davis (1907, p. 49) writes that "The adult Hyla andersonii
is amusingly active at night and jumps about the lower limbs of the trees and
on the bushes with much agility. They seem rarely to climb over five or six
feet from the ground. They sit upright and look pert, and if interrupted in
the midst of their song they leave their bubbles blown up until such a time as
the intruder goes away or stands still. In the day time they are usually quiet
and for the most part hide in the deep moss and leaves lying on the ground."

Miss M. C. Dickerson (1906, pp. 132, 133) had this species under observa-
tion in captivity and for a long time:

"The creature has a very gentle and alert expression. It is in fact, one
of the most alert and timid of our tree frogs. In captivity it is seldom content
until it finds some moist hiding-place in moss and ferns or under wet decaying
pieces of wood. When such a place is found, the Hyla backs into it with the
burrowing movements common among the Salientia. Here, with flattened
body and closed eyes, it remains sleeping away the days and nights. The
sleeping position is interesting, in that the long delicately-colored toes are
lifted to the green of the sides, the rest pressed close to the body just above
the shoulder.

"This Hyla is relatively sturdy. The tree frog found June i, 1888, was
still alive and well in January, 1889. How much longer it lived is not recorded.
The specimen caught in September, 1904, is in plump condition now in June,
1905.

"This specimen photographed moulted the skin February loth. For
several hours before the moult the skin was very dry and lustreless, and the
frog kept rubbing one hand and then the other over the head and eyes, as if
in discomfort. The skin was thin and white. It was shed in one piece and
swallowed as it was shed, in just the manner described for the similar process
for the toad and frog.

234 Frogs of the Okefinokee Swamp

"This Hyla differs conspicuously from Hyla versicolor and Hyla regilla,
in fact from most of its nearest relatives in North America, by the fact that
it has apparently but a limited power to change colour. The green may pale
or deepen, it may even take on a tinge of brown; the lavender tints of hand
and foot may be extremely light, or so dark that they become a deep purplish
brown. But when we compare this with the radical and rapid changes in
Hyla regilla and others, the limitation is very evident. As far as observed,
the pattern of dress is never obscured.

"Hyla andersonii is an agile chmber, but ascends largely by clasping the
hands and feet around the support. The adhesive power of the disks is
relatively not great, and the under surface of the body does not seem to aid
them as much in climbing as it does many tree frogs. On a relatively smooth
vertical surface up which Hyla cinerea, versicolor, pickeringii, regilla, and
many others will climb rapidly and with confidence, Anderson's Hyla holds,
itself with great difficulty, even for the space of a few seconds."

VOICE

The records of capture and voice characterizations make up much of the
literature of this species. The call has been and is the clue to its whereabouts.
Like each of the Hylas its voice is specific and distinct. But voice characteriza-
tions whether by syllables, figures of speech (similes, metaphores), musical
notation, human imitations or other means are dubious evidence from which
to draw conclusions. For example, John E. Peters (1889, pp. 58, 59) "On
June I, 1888 (he) found a specimen of Hyla andersonii Baird in a wet place
on the border of a pine barren at May's Landing, N. J. It was quite lively
when caught, but it soon became sluggish in confinement. Its voice was
shrill and light, comparatively speaking; and it consisted of a repetition of
the smae note three or four times in regular succession, in a sort of 'peep, peep,
peep, peep,' as nearly as I can give it. The specimen was sent alive to Dr.
C. C. Abbott, of Trenton, N. J. who says in his 'Catalogue of the Vertebrate
Animals of New Jersey' (Geology of N. J., Cook 1868, p. 805) that it is 'a
Southern species, a single specimen of which was found in Camden County
in 1863' by Dr. J. Leidy. . . . The specimen is still alive (Jan. 1889), and may
be seen by applying to George Pine, Esq., Trenton, N. J."

A year later the Dr. C. C. Abbott above (1890, p. 189) wrote as follows:
"The specimen of this beautiful batrachian referred to by Dr. Peters in the
January NaturaUst is still in excellent health (Feb. 1890), and occasionally
utters its characteristic cry, which should not be described by the word
'peep' for it suggests a similarity to the cry of the Pickering's Hyla, which
shrilly 'peeps'. The andersonii utters a single note, better described by the
syllable 'keck', which it usually repeats three or four times. It is not a frog-
like note at all, but much resembles the call of the Virginia rail (Rallus
virginianus). If the collector follows up any 'peeper' in the marshes, he will
not discover additional specimens of Hyla andersonii."

Of the same individual H. W. Fowler (1908, p. iii) writes "Dr. Abbott
tells me that the example Peters sent him from May's Landing alive uttered

Hyla andersonii 235

a sound more like a clatter, or sharp click, something like that made by the
Virginia rail. The vocal vesicle resembled that of Hyla versicolor. It appeared
to take in a big breath and produced about 6 clicks as the air escaped."
Thus it appears the doctors (M. D.) do not agree. No one reports how
George Pine might have characterized this individual frog. Furthermore,
there are many people who have characterized in my presence Hyla
crucifer, Pseudacris triseriata, Pseudacris nigrita, Acris gryllus as "peep"ers
although the writer might not do the same. If one used the "keck" character-
ization of Abbott and Harper's kek for Hyla femoralis one might infer close
relationship. The latter, however, characterizes the call of Hyla andersonii
as "quak". The calls are different as both Harper and I know from experience.
From the diverse characterizations of the call of Hyla andersonii Noble and
Noble (1927, p. 426) write "One would gather from the literature that' the
Anderson Tree Frog had a variety of calls", but they proceed to state it has
one call thereby implying quite rightly that the different renditions of its
call are for one call not many. But the abnormal calls of the tree toad re-
corded by Wright are assigned by them to Overton's "turkey root" call.
One call is that call but others are not. Herein is revealed the difficulty of
one person interpreting the call evidence of another. Overton, Harper and
the writer have been abroad at night together, so by direct identification we
know what the other means by each syllabic or simile characterization, but
I would hate to have to identify a frog without this aid though hearing is one
of my best means of identification. No two have characterized more frogs
nor better than Overton and Harper, yet their characterizations are not the
way I always hear them.

In 1894, J. Percy Moore (1894, pp. 1045, 1046) writes of its voice and
habits thus: "About the middle of June, 1889, Mr. Louis M. Glackens and
the writer were engaged in general biological studies along the Atsion and
Batsta Creeks in Atlantic and Burlington Counties, New Jersey. On the
night of June 17th we stopped at Pleasant Mills. Shortly before sundown a
thunder storm arose, just previous to and during which the frogs became very
noisy in a swampy thicket near by.

"The note was an unfamiliar one and invited investigation, which resulted
in the capture of two specimens of this handsome and rare species. The shrill
quack-ack, which at the time was compared to the note of a frightened guinea
fowl, and which is not unlike the call of a rail, was constant and seemed to
come from every tree, but during our progress through the voices immediate-
ly around us, for a radius of about 25 feet, were silent. This circumstance,
and the oncoming darkness made it difficult to secure specimens, although
frogs were so abundant. The two secured were found perched on the lower
side of branches of pines with dilated and vibrating throats, though at the
moment they were silent; and it was noted that they emitted an odor which
was likened to that of raw green peas. . . . The following morning we could
find no trace of them, but later in the day heard another chorus in the middle
of a dense swampy thicket. Since then Mr. H. F. Moore and myself have

236 Frogs of the Okefinokee Swamp

repeatedly visited the locality in quest of the Hyla and its eggs, but entirely
without success."

W. T. Davis who has done more than any other individual to introduce
this species to naturalists writes of it in 1905 (p. 795) as follows: "At Lake-
hurst, New Jersey, especially in the latter part of May and in June, there are
heard at evening coming from the white cedar swamps, many voices that
resemble the familiar quacking of ducks. If one will take the trouble to wade
into one of these swamps at twilight and approach the singer cautiously, it
will be discovered that he is a male Hyla andersonii. He pipes and sings
'aquack-aquack-aquack' many times, or until his bubble of air gives out.
This is the time, when he is singing, to take a step forward. Even when the
observer is very near and evidently plainly in view, he cannot resist the
temptation to sing, for he hears his rivals all about calling loudly. The notes
are not all alike in sound, and some individuals remind one of the 'potrack-
potrack' of the farm and guinea fowl. The Hyla will be found seated on the
lower limb of some tree, or among the top branches of a huckeberry bush. I
have heard this frog singing at mid-day when the sun was shining brightly,
especially after a shower. Also solitary individuals may be heard in the swamps
much later in the year, and they do not appear to wander as far from the
water as does Hyla versicolor''. In 1907 (1907, pp. 49, 50) Davis recorded
them calling in August. "On the warm cloudy evening of August loth Hyla
andersonii was heard near some pools a short distance north of the village of
Farmingdale, N. J."

In 19 16 T. Barbour (19 16 pp. 6, 7) and his brother F. K. Barbour, of their
observations speak thus: "At dusk we have usually taken our supper, and
then waited for darkness to come on and for the Hylas to begin to sing, and
have had good luck taking Hylas by the following method : One of us with an
electric flashlight would start for the nearest singing Hyla, while the other
usually waited some distance away. As soon as the Hyla stopped singing, the
person who was not trying to approach would imitate the call of the frog, and
this would start it singing again vigorously, and while it was singing the
collector bearing the light would approach as quickly as possible, standing
still as soon as the singing ceased. This process was kept up until finally the
light flashed on the vibrating white throat of the singing Hyla, and its capture
then became a perfectly simple matter, as long as they stared stupidly at the
brilliant light.

". . .This year (1915), however, we did not get down to Lakehurst until
the 8th of July, when we found the Hylas singing in goodly numbers in the
white cedars about the lake. ... A large number of individuals kept on
singing so vigorously until at least the 20th of July. I might add that the
singing begins with dark, though an occasional voice may be heard upon a
lowery afternoon, and the greatest concert takes place before ten o'clock. At
about this time, the number of singing individuals is noticeably less."

Thus from 1889-1918 the "calling" season appears from records to be
from May to August 10.

Hyla under sonii 237

Noble and Noble (1923, pp. 426, 429) recorded that: "//. andersonii has
only one call and that is given with fully inflated pouch. It is a series of ten
to twenty, or even more, resonant nasal quanks. If one tries to shout the
word quank while holding the nostrils closed, a sound is produced not unlike
the note made by this frog. The call sounds somewhat different from a dis-
tance, especially when several frogs are calling at once. Then the notes tend
to run together, each note having two syllables, a-quank, a-quank".
"i. H. andersonii begins calling in early May from concealment on the
ground.

2. In the middle of May and throughout June and July the males call
chiefly from bushes or from trees.

3. At various intervals throughout May and June (and some years in July)
when the rains have flooded the bogs and changed the ditches into small
sphagnum-choked streams, the males leave their calling stations and make
their way to the nearest of these small streams. This migration occurs
about midnight. The males begin calling again from their new positions
near the sphagnaceous streams. ..."

They heard Anderson tree frogs from May 13-14 to July 23.

On June 8, 1922, on our way to Okefinokee Swamp we camped in the
evening on the north side of Everett's Pond, N. C. near the South CaroHna-
North Carolina line. The instant it became dark "We heard plenty of Acris
gryllus, Bufo fowleri, Rana catesheiana, a few Rana clamitans, Hyla cinerea in
chorus, Hyla versicolor, quite a few. In the wooded sections or streams
following into the lake in dense thickets, S7nilax-briar entwined, were Rana
virgatipes croaking quite general. Fine! Several in different places. Later
Harper went above his tent and heard a strange note some 20 rods away.
The note was lower than that of Hyla cinerea, much more rapid than it,
possibly like it in form, yet not. In some ways it seems quite different from
Hyla cinerea. Captured this one nearest the tent. Another individual
heard near a negro cabin. We could not get it because we strangers did not
wish to disturb them. We heard another near the cabin. This at a distance
sounded quite different from Hyla cinerea down by the lake.

Later in the evening Miles Pirnie heard one opposite our camp. All the
frogs heard were in or near the lake. These H. andersonii were in a stream
or branch with sweet gum, tangle of oaks, bamboo briars (Stnilax), Magnolia
glavca, maple, black gum and red maple.

Harper speaks of the calling of these Hylas thus: "I heard the note of
H. andersonii at a distance of 200 yards and suspected almost at once what .
it was. Trailed it and after a long wait located it in some tall bushes in the
edge of a branch swamp. Its note bears a general resemblance to that of
H. cinerea but goes about twice as rapidly, is about half as loud and sounds
more like quak, quak, then quonk. It carried fairly well at 200 yards and
about 300 yards would probably be the limit. . . . Did not see it croak. Its
periods are infrequent and irregular, 2 minutes interval. Perhaps 15 or 20
or 25 notes given in one period."

238 Frogs of the Okefinokee Swamp

On June 28-29, Mrs. Wright and I heard these Hyla andersonii in the
wooded edge of Mr. P. H. Emhe's lake at Lakehurst, N. J. Some of the
principal trees and shrubs where the males were numerous were: Magnolia
virginiana, Acer ruhrum, Vaccinium corymhosum, Vaccinium atrococcwm,
Azalea viscosa, Chamaecy parts thuyoides, Nyssa sylvatica, Ilex glabra, Clethra
alnifoliutn, Vihurnum nudum and Rhus vernix. Again on May 23, 1924,
we heard a few, though as on the former occasion our attention was mainly
directed to Rana virgatipes tadpoles and transformed frogs.

In June 1928 (i5th-28th) Alexander Barrett Klots and Elsie Brought on
Klots made these detailed notes on several frogs found at Weymouth in
Atlantic Co., N. J.

"Many attempts have been made to reproduce the quality of the song
in print, with results differing for practically every observer. Thus the call
has been variously recorded as "peep," "keck," "quak" (Harper), "quack-
ack" hke a frightened guinea fowl, and "quank." To the writer the call
seemed a nasal "quack," almost verging on a "quank" but without the strong
"n" sound of the latter. The call was never disyllabic.

"The note is repeated at about half -second intervals for sometimes fully
30 seconds. When the frogs are in full song an interval of about two minutes
intervenes between outbursts. We had no difficulty in starting the frogs
calling again at distances of from fifty to three feet, after they had been silent
for a minute of two. One individual was recorded as having called 74 times
in one period of song.

"The frogs definitely associate together for singing, whether because of the
presence of females or for companionship. The latter probably plays a con-
siderable share in the performance, as is evidenced by the quick response to
an imitation of the call. Five such singing groups were definitely located.
Of these the first contained seven individuals, the second contained three,
the third contained eight, the fourth contained three and the fifth, which was
just across an uncrossable creek, contained at least six. Only once was a
single individual noted in song alone, and that was a frog which called three
times in a spot a half-mile distant from any others and was never heard from
again.

"The locations of the groups were fixed, and during our stay did not change
a particle. Night after night a group would be in exactly the same area,
though the individuals composing it shifted position a bit.

"The time of singing was remarkably constant. On every night but one
the chorus started between ten and fifteen minutes before sundown. On the
one exception, a clear dry night with a bright moon, the first songs were not
heard until twenty minutes after sundown.

"The carrying power of the song was excellent. A chorus was plainly
heard as an entity over 800 paces away, with two patches of woods and a
brushy swamp intervening. The wind was negligible. Individual voices were
distinguishable 754 paces away down a straight road, with a light wind blow-
ing from the observers toward the frogs."

Plate XX\'
Anderson tree-frog {Hyla andersonii)

1. Mr. Emlie's field, Lakehurst, N. J. June 28, 1923.

2. Male, Everett's pond, N. C. June 9, 1922. Ventral aspect, x i.o.

3. Male, Everett's pond, N. C. June 9, 1922. Lateral aspect, x i.o.

4. Tadpoles, Lakehurst, X. J. June 28, 1923. Lateral aspect, x 1.0.

5. 6. 7. 8. 9. Series from tadpole to transformed frog. Lakehurst, N. J. July

3, 1923. X I.e.

^^^^

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S-^f ■ ■--! . ' /*-^^f"^A?^'

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f f 4

Hyla andersonii 239

MATING

Male. — {From life). Upper parts (dorsum, upper lip, angle of mouth and
dorsal surfaces of limbs) cress green to a light cress green, in sides lighter to
deep chrysohte green. Stripe along side, behind vent, along limbs, upper
jaw cartridge buff, ivory yellow or marguerite yellow or seafoam yellow. Area
back of and along side of vent vinaceous drab, purple drab becoming over
tympanum dark purplish drab. A little of same color scattered on throat.
Tops of forefeet and hind feet deep brownish drab except for first two digits
which have the cadmium yellow spots on the top. Fore part of under side of
antebrachium and under part of brachium with cadmium yellow — deep
chrome or orange spots, also a little area in axilla ; also groin and most of fore
and hind part of femur, whole undersides of tibia and inner side of foot. All
these orange or deep chrome spots on a raw sienna or mars yellow background.
Iris more or less vinaceous drab or purple drab with vinaceous tawny spots.
Under parts white except for dark purplish drab of throat.

Female. — {From life). Throat white, grayish or with slight purplish drab
tinge. Green patch below angle of mouth usually white edged, usually absent
in males.

Structural differences. The differences in color already noted, and larger
females than males are the perceptible differences. Noble and Noble (1923,
p. 433) note that the male has minute asperities on inner and upper side of
thumb. They gave average length from vent to snout of ten females as 40.9
mm. (extremes 44 and 38 mm.), of fifteen breeding males as 36.7 mm. (ex-
tremes 35-38 mm.). The females, as in most of our Hylas, are larger. Al-
though some authors have spoken of them as "about 2 inches in body length, I
have seen none over 47 mm. in length. The range of males seems to be from
30 mm.-4i mm. in body length; of females 38-47 mm. No doubt sexual
males below 30 mm. and females somewhat below 38 mm. will be found.

Duration, night or day. Amplexation. All the records of mating by Bar-
bour, Davis and Noble and Noble were for the evening. In each case ap-
parently the pair broke after laying. In no instance are abnormal embraces
recorded or mentioned. All doubtless are axillary or supra-axillary. The last
authors speak of "a normal supra-axillary amplexus" and describe the
amplexus.

OVULATION

Habitat and period. Barbour (19 16, pp. 6, 7) the first to record a mated
pair and secure eggs writes "Up to this year (1915) I have known nothing
regarding the breeding habits or the time of egg-laying of these Hylas, but
have assumed that it was in May, as Davis reported them singing very plen-
tifully at that time. This year, however, we did not get down to Lakehrust
until the 8th of July, when we found the Hylas in goodly numbers in the white
cedars about the lake. After capturing a number of singing males (I had
never taken a female before), my light flashed by the merest chance upon a
pair of Hylas sitting well up in a pine tree, in embrace. This, and another
taken, in a similar situation, were the only females secured, although we took

240 Frogs of the Okefinokee Swamp

several males from the low oak scrub about a small fresh water pool in the
pine barrens. All of our catch was placed in a large mouthed jar and brought
home to Rumson alive. The next morning a number of gelatinous egg-masses
were found in the bottom of the jar. Since they were pretty well smashed
up from the struggles which the Hylas made trying to escape, it was impossible
to distinguish whether the egg-masses were laid in a characteristic form. We
were both surprised to find these eggs, as we had not supposed that this Hyla
laid so late in the year; yet we had always wondered why such a large number
of individuals kept on singing so vigorously until at least the 20th of July."

Davis and Chapin secured ovulation June 13 and 14, 1922. Noble and
Noble recorded ovulation May 20-21 and June 4-5, 1922. For the species,
ovulation records then obtain for May 20-21, 1922, June 4-5, 1922, June 13-
14, 1922 and July 8, 1915, or from May 20 to July 8. If a few tadpoles
transform in late June, as we observed in 1923 at Lakehurst, possibly breeding
may some years begin as early as May i. Doubtless with this species, as
with most frogs and certainly southeastern United States frogs, humidity
(rains) is the important external factor in onset of breeding. In this species,
as in all our southern frogs, it is a long season breeder or protracted breeder as
Noble has called it. But exposure and protracted breeding classifications are
local at the most, e.g., Sca'phiopus holbrookii and all Scaphiopi may be very
protracted and generally are. Rana pipiens may or may not be long extended
in breeding season. Most southern and southwestern forms have long breed-
ing seasons.

Thus far the eggs laid in nature have been laid in sphagnaceous streams or
puddles near by. Once upon a time we were ready to call Rana virgatipes a
Sphagnum Frog on the basis of Lakehurst experience but it is not wholly so.
So, too, H. andersonii may be more widespread in its place of ovulation. One
wonders whether in Cheraw, Anderson and Southern Pines it is limited to
sphagnum. We did not casually record sphagnum at Cheraw but we did not
search for it. In a similar way we suspected they may reach higher than 6-8
feet in the trees when not ovulating.

Egg-laying process. Noble and Noble (1923, pp. 433-439) gave detailed
attention to the interesting egg oviposition, making observations on two pairs
in the field and two in the laboratory. It is too explicit for me to paraphrase
and too extended to quote in toto.

EGGS

Noble and Noble (1923, p. 439, 440) distinguish the eggs thus:
"i. Single, not adhering to one another, usually scattered among the water
weed.

2. Attached to sphagnum (rarely debris) or free and rest on bottom.

3. Found on bottom of small, non-stagnant pools, or in slow-moving streams
of the pine barrens.

Before cleavage the cap of the animal pole is usually dark brown, the
other two-thirds of the egg, creamy white. . . ." Hyla andersonii as they dis-
cover like many other North American frog's eggs change in pigmentation

Hyla andersonii 241

soon after they are laid. To continue with the description. "The egg is
surrounded by the vitelhne membrane and by the two gelatinous membranes
of the usual type. The gelatinous capsules vary enormously according to
their age and treatment. The following measurements are taken from a
series preserved in formalin. They agree well in size with some living speci-
mens. Diameter of Ovum — 1.2-1.4 mm. Diameter of Inner Capsule —
1,9-2.0 mm. Diameter of Outer Capsule — 3.5-4.0 mm. The vitelline mem-
brane may be best demonstrated after maturation when the animal pole is
slightly flattened leaving a space between membrane and ovum.

"In nature, H. andersonii was estimated to lay eight hundred to one
thousand eggs. None of our laboratory animals laid more than eight hun-
dred eggs."

HATCHING PERIOD

Noble and Noble (1923, p. 441) had eggs hatch in laboratory in four days
and beHeved it would take longer in nature. In South Carolina or North
Carolina it might even happen in less than four days. In New Jersey it
might be longer or shorter.

MATUEE TADPOLE

Color description from life {May 23, 1924)- General coloration of body
olive, brownish olive, or dark olive. Back with scattered black spots. Gill
region with a sheen of ochraceous salmon or vinaceous tawny or vinaceous
salmon. Interspersed with this color is some black. The mental region is
clear of color. Below eye blotched with black, bronze (or one of three gill
region colors) and green yellow. Over the belly in younger specimens is a
full block of ochraceous salmon or vinaceous cinnamon. In older specimens
green-yellow or citron yellow or sometimes seafoam yellow across the belly
or on the gill region.

Tail. Starting from the base of the tail there is a longitudinal irregular
blackish band along the musculature of the tail one-half of an inch or more
from the tip of the tail. This band is more or less interrupted, not a clearly
defined band as in Hylafemoralis or in Pseudacris ocularis or other Pseudacris
species. Below the band is a clear area of warm buff or cream color or car-
tridge buff or seafoam yellow. In some specimens there is a similarly colored
area 1/8-1/4 inch long above the dark band. The very lower edge of the
musculature is with a fine purplish line. Just above this edge are a few scat-
tered collections of black dots. Upper and lower crests heavily clouded with
blackish dots which in places assemble in clusters. In younger tadpoles quite
a prominent irregular margin of blackish on the crests and the musculature
band of black more regular and not so interrupted.

Eye with pupil rim prominent ochraceous salmon or vinaceous tawny
or vinaceous cinnamon. Rest of pupil spotted with this color and black.
All in all it is a prominently colored eye.

General appearance. Tadpole (35 mm.) full and deep-bodied. Tail
medium to fairly long, tail tip acuminate or sharply acute. The dorsal and
ventral crests about equal. The dorsal crest is less than the depth of the

242 Frogs of the Okefinokee Swamp

musculature and extends onto the body to the vertical midway between hind
legs and the spiracle. Spiracle sinistral directed upwards and backwards,
far below lateral axis, the spiracular opening plainly visible and elliptical or
round. Eye just touches or is below the lateral axis, is in dorsal aspect on the
lateral outline and in consequence visible from the venter. Anus dextral,
opening at or only slightly above the lower edge of the ventral crest. Mu-
ciferous crypts not distinct.

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
upper fringe for about 2/7-1/4 of the length of the upper fringe. The end
of the second upper labial row may be even or beyond the end of the fringe.
The horny beak is contained in upper fringe 1.5-1.7 times. Median space
between the lateral second upper labial row 1.25-2 in the length of either
lateral row. The lateral row is contained about 2 1/2 in the upper fringe.
The inner papillae extend beneath the third lower labial row of teeth giving a
two-rowed appearance of papiUae below it hke Hyla cinerea. The third row of
labial teeth is short (in one specimen subdivided) and is 2/9-1.3 times
(usually 2/5, rarely as small as 1/8) in the length of the first or second rows.
The first and second rows about equal or i 1/2 greater than the horny beaks.

Measureryients. Length of body (10. 0-12.0 mm.) in tail (i 1.5-23.0 mm.)
1. 1-2.0, average 1.67. Width (6.0-7.2 mm.) of body in its own length 1.5-
1.85, average 1.65. Depth (5.0-6,5 mm.) of body i. 0-1.85 mm- in body
width, average 1.2. Depth of body i. 61-2. 18 in body length, average 1.86.
Depth (5.0-7.0 mm.) of tail in length of tail 2.5-3.5, average 3.0. Muscular
part (2.0-3.0 mm.) 1.9-2.5 in depth of tail, average 2.1. Spiracle i. 0-1.5
nearer base of hind legs or vent region (5.0-7.0 mm.) than the tip of the
snout (6.0-9.0 mm.), average 1.22. Spiracle 1.25-2.3 nearer eye (2.5-4.8
mm.) than base of hind legs or vent, average 1.86. Eye i. 0-1.6 nearer to
spiracle (2.5-3.5 mm.) than to tip of snout (3.0-4.2 mm.), average 1.25.
Nostril 1. 2-2. 1 nearer eye (1.2-2.5 mm.) than snout (2.5-3.0 mm.), average
1.5. Internasal space (2.2-3.2 mm.) usually .75-1.6 larger than mouth
(2.0-3.0 mm), average 1.25. Mouth contained 1.33-2.5 (average 1.85) in
interorbital distance (4.0-5.5 mm.). Internasal space contained in inter-
orbital distance 1.33-2.2, average 1.8.

The dimensions of the largest tadpole are:

mm.

mm.

Total length

35

Spiracle to vent

6.0

Body length

12

Spiracle to eye

4.8

Body depth

5-5

Eye to snout

4.2

Body width

7.2

Eye to nostril

2-5

Tail length

23.0

Nostril to snout

3-0

Tail depth

6.3

Mouth

3-0

Musculature of tail

3-0

Interorbital distance

S-o

Spiracle to snout

9.0

Internasal distance

2.8

Hyla andersonii 243

General remarks. The first author to discover and to describe the tad-
poles is W. T. Davis (1907, pp. 49, 50). "It was observed in June at Lake-
hurst that the males of Hyla andersonii were attracted to a few small pools
in particular, several of which were only a yard or two in diameter. On July
2ist, with Mr. James Chapin, I made search of these pools for the tadpoles,
and was fortunate in finding a number in one pool, although they appeared
to be absent from another and similar locality about a mile distant where the
adult frogs had been and were still most numerous. The tadpoles collected
were in all stages from a few millimeters long to those just leaving the water
as little frogs. The mature tadpoles are from 35 to 40 mm. long and of the
usual tadpole color, that is, of the color of the muddy bottom of a pool. The
under parts are lighter and show a golden sheen, which sometimes extends
up the sides. The small hind legs show early on the ends of the toes the disks
that are so conspicuous in' the mature Hyla. The tail is spotted, and there is
usually a dark irregular marginal band. The maculations sometimes become
irregular blotches as on the tails of the tadpoles of Hyla versicolor."

Noble and Noble (1923, pp. 443-448) give a detailed account of the tad-
pole of this species with illustrations of labial mouth parts and the mature
tadpole.

LARVAL PERIOD

We found the larvae in several pools, grassy, sedgy, sphagnaceous along
a dense woody border below Mr. P. H. Emlie's lake at Lakehurst. The follow-
ing trees and shrubs were in the wooded edge where numerous H. andersonii
were calling and taken. The tadpoles also were in the more or less open
shrubby areas where Voccinium and Magnolia (small) were common. A few
pools were connected with streamlets and water ditches.

The tadpoles we took June 29 began transforming July 2. This species
actively croaks in May in other years. These tadpoles doubtless were at
least from eggs hatched in the first half or middle of May. My determination
is from comparative experience with Hyla squirella, Hyla femoralis, Hyla
cinerea, Hyla gratiosa, Hyla arenicolor and Hyla crucifer. At the time
at Lakehurst we concluded 50-75 days a probable larval period. Noble and
Noble found transformation July 23. If these transforming frogs were from
eggs of May 20 or June 4, we have 49-64 days for larval period. Transforming
frogs of July i and 2 apparently from earlier seasonal oviposition might take
10 or more days longer.

TRANSFORMATION

Period. Messrs. Davis and Chapin found this species transforming from
tadpoles to frogs July 21, 1906 (Davis, 1907, pp. 49, 50). On June 29 the
author and Mrs. Wright found a few tadpoles with two legs well developed
and by July 2-4 they began to transform and travel about my screened porch.
The last transformed July 15. The bulk of these tadpoles at Lakehurst must
have transformed by July 15-30. Noble and Noble found them transformed
July 23, 1923. The eggs taken by Barbour (1916, pp. 6, 7) JulJ^ 8 doubtless
would not have changed to froglets until August 25 or September i. Judging

244 Frogs of the Okefinokee Swamp

from choruses we heard June 26-30, 1923, and previous observations by others
the transformation doubtless extended from June 29-September i with
the majority changed into frogs before August i.

Size. "When the tail is nearly absorbed, and they leave the water, they
are about 25 mm. long and of a dull olive green. They grow lighter, that is,
brighter green in hue with the disappearance of the tail until the little frogs
which in length of body are 15 mm., resemble the mature individuals. The
white that margins the green of the back and extremities is not so conspicuous
as in the adults, and the saffron of the under parts is wanting in those that I
have examined." (Davis, 1907, p. 50).

In color the transformed frog looks much like that of Hyla femoralis or of
Hyla squirella. The dark vitta does not extend along the side but is a pur-
plish area before and back of the eye just before the tail is entirely absorbed.
Not until the four legs were acquired did our froglets have green appearing
on the back.

The range in body length of one collection of 10 individuals is 11. 5- 13.0
mm.; average 12.0 mm.; mode 12.0 mm. Our transformation sizes, therefore,
prove slightly smaller than those of Mr. Davis. The two records combined
give a range of 11-15 mm. for transformation size.

GROWTH

This species begins as little froglets at 11. 5-15 mm. in body length. Dr.
Barbour has material, 13 specimens (M. C. L. Nos. 3218, 3577-79, 3 55 1-60)
from Lakehurst most of which was taken by himself and F. K. Barbour.
These measure 35 d^, 36 d", 36 cf , 37 cT, 37 d", 37 d", 38 d", 38 d", 38 d",
38 cT, 38.5 d^ mm., 38.5 d^, 40.5 d^ ; and one female 44 d^ mm. This gives a
range of males from 35-40.5 mm. The Nobles (A. M. N. H. Nos. 16960-83)
had males 35-38 mm. and females 38-44 mm. From comparative experience
they may prove 3 years old, but our collections and collections in general
have so little material between 15 mm. and 35 mm. that it is useless to attempt
an approximation of this question. Possibly some adults of second year from
26 or 27-33 or 34 mm. may breed.

FOOD

Noble and Noble (1923, pp. 448, 449) record that "The stomachs of ten
males which were captured during June while caUing from bushes or low
trees contained the following food: 5 grasshoppers (two species); 2 beetles,
3 ants (two species), i dipterous insect, 2 dipterous pupae (tabaniid?), and
some unidentifiable insect remains. None of the specimens taken in embrace
contained food in their stomachs, but only a few pairs were killed immediately
after oviposition." Dr. Burt Green Wilder made extended notes on food and
food prehension in this species. He kept a male (sent by Dr. James P.
Chapin) from July 21, 1906 to November 21, 190,8, when the creature was
killed. It, therefore, lived in captivity over two years. Unfortunately his
notes are not to be found. This is the same specimen L. A. Fuertes painted.

Hyla andersonii 245

ENEMIES

There are plenty of water snakes in the neighborhood which might prey-
on the adults when in ponds, and on journeys to and from breeding places.
In some localities the associated Chrysemys pida, Clemmys guttata, Chelydra
serpentina and other turtles do damage to tadpoles. In general the associated
fish, such as Acantharchus, Enneacanthus, Mesogonistius and Umbra doubt-
lessly prove of little consequence in the economy of the tadpoles. The eggs
are laid in shallow places where the fish are not so common.

AUTUMNAL DISAPPEARANCE

The last date of appearance is September 5, 1904, (Davis, 1904, p. 893).
"Mr. J. A. G. Rehn has reports of one seen August i6th, 1907, at Stafford's
Forge, in Ocean County. It was probably driven from the woods by a forest
fire" (Fowler, 1908, p. 193). Of course, it must retire into winter retreat much
later.

AFFINITIES

Cope likened it to Hyla arhorea as did Boulenger (1882).

Cope (1862, p. 155) in speaking of the first specimen he ever saw alive
and from which this color description from life comes writes that,

"I am indebted to Dr. Jos. Leidy for a beautiful specimen of this frog.
It was found in a cedar swamp, near the town of Jackson, in New Jersey,
sixteen miles east of Philadelphia. Without careful examination of the
specimen, he supposed it to be the viridis of the Southern States, from its
great resemblance to that species, and presented it as such, at the meeting of the
Academy the same evening (Vid. Proc. Acad, for July, p. 305). At the same
time Baltimore was given as its northern limit upon the authority of Dr.
Uhler of that place. As Dr. Holbrook gives lat. 33° as the most northern
habitat known to him, it would be interesting to receive specimens from
Baltimore, as there is a possibility of the supposed viridis being the andersonii.'

At this time Cope (1862, p. 154) and later (1889, p. 365) likened it to
Hyla arhorea as did Boulenger (1882).

Barbour (1914, p. 239) has compared it with Hyla pulchrilineata (Santa
Domingo) the form Boulenger (1882) places just before Hyla andersonii in
his Catalogue. Noble & Noble (1923) assert that "Hyla andersonii is not
closely related to H. arhorea nor to H. pulchrilineata.'' "The relationships of
Hyla andersonii are to be sought in Chinese forms and probably in H.
immaculata Bott (Boettger)". It may not be related to Hyla arhorea. It
is interesting to attempt to hnk it with a Chinese form, but somehow the
author is incHned to believe we must search for an American forebear. " Hyla
andersonii has been derived from a group of Hylas which laid surface eggs,
its method of oviposition being a modification of their method." The present
writer who has seen in nature and laboratory as well the eggs of almost all
the Hylids of the U. S. A. would not unhesitantly say this is the case though
he is inchned toward it.

246 Frogs of the Okefinokee Swamp

In general coloration it falls in the group of Hyla cinerea, Hyla cinerea
evittata, Hyla gratiosa (sometimes clear green), Hyla squirella (sometimes clear
green). Several of these species in males may have green from fore arm for-
ward on either side of the throat. In some individuals almost to symphysis
of the jaw. In Hyla andersonii it is more definite (white edged usually in the
female). (See extended discussion under Variation in color Hyla cinerea). In
general, the upper parts of Hyla andersonii are uniform. Sometimes though
numerous Hyla cinerea without white or yellow dots or spots on back may be
collected but usually they have them. Hyla gratiosa usually has its character-
istic dorsal spots but occasionally it may be green with a few dorsal yellow or
white spots. Am sure I saw one Hyla andersonii male with a dorsal yellow
spot or so but haven't the specimen now to prove it. Too great emphasis
cannot be laid on any one character of coloration in Hylas in estabUshing or
eliminating relationships. LeConte (1825) questioned Daudin's green Hyla
squirella description and illustration but green they will become at times.
The above Hyla species are also sexually dimorphic as well as Hyla andersonii.

If Hyla cinerea with lip and lateral white or yellow stripe with thin
purplish line below appear with light line different in extent, purple below
absent or present, light line on upper lip only, or without lateral stripe at all
{evittata) it can also intensify the purplish below the white hne to get the
effect in Hyla andersonii. In a somewhat similar way Hyla squirella and
Hyla andersonii might be linked hypothetically. It is, however, hypothesis
and is so involved the writer handles it warily. Voice is uncertain in evalua-
tion of relationships. Each person might hear it differently, use musical
notation, syllables, lines and other forms of characterizations. Too many
personal equations are involved unless one person heard all forms to be con-
sidered. In egg deposition its eggs are single placed (several emitted at one
time) but more or less singly placed in dropping to the bottom. Its method
of laying eggs is very interesting. This might lead one to suspect that when
some local naturalists of the South can devote some time to Hyla squirella,
Hyla gratiosa and other species some of these may have a similar method of
deposition. To be certain it must be field and laboratory determined and
well checked which most itinerant naturafists have not the time to do thor-
oughly. Noble and Noble describe the egg minutely. Without the egg in
hand, the writer hardly can say whether it is of Hyla squirella affinity (two
envelopes) or Hyla gratiosa (single envelope) or Hyla cinerea (two envelopes).
In some way Hyla cinerea with both surface film and strewn eggs in submerged
bladderworts, etc., may be the nearer form though Hyla gratiosa and Hijla
squirella have singly strewn eggs.

In general coloration the tadpoles look like Hyla femoralis. The first
tadpoles of Hyla andersonii that the writer saw at Lakehurst reminded him
very decidedly of Hyla femoralis. Knowing this form was not in Lakehurst
we naturally placed them in Hyla andersonii. By accident they proved the
very pools in which Noble and Noble worked so hard the previous year. Super-
ficially I took them to be near Hyla femoralis. Closer inspection made the
resemblance less apparent. They do not superficially and clearly fall into the

Hyla andersonii 247

group with heavily spotted and somewhat scarlet suffused tails ( H. arenicolor,
H. femoralis, H. versicolor). Neither do they superficially fall into the
greenish tadpoles of H. gratiosa, Hyla cinerea group. Examinations of their
mouth parts apparently put them near this group (short third lower labial
tooth row and other characters).

This topic could be expanded at will. The value of certain criteria in
determining its possible relationship to Hyla arhorea, or to Hyla pulchrilineata
are not for me to discuss. If we let license run riot we might call it a northern
representative of H. gratiosa, or an offshoot from Hyla squirella as a radiation
center or a Hyla cinerea derivative just as Hyla c. evittata apparently is. The
writer is still searching for slow coming material before he indulges in that
joyous privilege of final pronouncements on questions of relationships.

Though not exactly pertinent to the topic of affinities possibly the follow-
notes from hfe may be interesting. In camp 1922 at Chesser Island we had
a male Hyla andersonii (from North CaroHna) and males of Hyla cinerea,
Hyla femoralis, Hyla versicolor d^nd Hyla squirella. Our rough journal notes
reveal the following notes made at one sitting under similar conditions with
fresh material.

Back. Hyla andersonii, cress green; Hyla cinerea, rear greenish-
yellow; head viridine; Hyla femoralis, brownish drab. Hyla versicolor, rear
pale smoke gray; rest grayish olive to vetiver green; Hyla squirella, buffy
brown or isabella color or old gold ; rear, yellowish citrine or apple green.

Degree of white on venter. Hyla andersonii, whitish on venter; Hyla
cinerea; Hyla versicolor; Hyla femoralis; Hyla squirella, least whitish on
venter.

Degree of orange. Hyla squirella, most orange below; throat, under side
of fore feet, fore limbs, hind limbs and hind feet fore and aft. Hyla andersonii,
front and rear of hind limbs when revealed ; under side of foot ; Hyla versicolor,
groin; front and rear of femur shows it slightly.

Green on throat of males: Hyla andersonii, Hyla cinerea, (Hyla gratiosa).

Iris. Somewhat of same color as back: Hyla cinerea, Hyla versicolor,
Hyla squirella, Hyla femoralis. Somewhat like vitta : Hyla andersonii.

Vitta. Hyla cinerea, Hyla versicolor, no vitta nor stripe through the eye;
Hyla femoralis, Hyla squirella, line in front of eye obscure or absent; Hyla
andersonii, vitta prominent.

BIBLIOGRAPHY
1868 Abbott, C. C. Geologv of New Jersey by G. H. Cook. Appendix E, p. 805.

1890 . The American Naturalist February 1890, Vol. XXIV, No. 278, p. 189.

1854 Baird, S. F. Proc. Acad. Nat. Sci. Phila., April 1854, PP- 60, 61.
1916 Barbour, T. Copeia, Jan. 24, 1916, No. 26, pp. 6, 7.
1907 Brimley, C. S. Journ. Elisha Mitchell So. XXIII, p. 158.

1863 Cope, E. D. Proc. Acad. Nat. Science Phila. Vol. 14, March 25, 1862, pp. 154, 155.
1887 Cope, E. D. The Batrachia of North America, U. S. Nat. Mus. Bull. 32. Washing-
ton, 1887, p. 365.
1904 Davis, W. T. Proc. Nat. Sci. Assoc, Staten Island, Oct. 15, 1904, Vol. IX, No. 8,
p. 26.

1904 . American Naturalist Nov. -Dec, 1904, Vol. XXXVIII, p. 893.

1905 • American Naturahst Nov. 1905, Vol. XXXIX, No. 467, p. 765.

1907 American Naturalist Jan. 1907, Vol. XLI, No. 481, pp. 49, 50.

1908 . Proc. Staten Island Assoc. Arts and Sciences, Feb.-May 1908, Vol. II, Part

II, pp. 49-50.

248 Fi'ogs of the Okefinokee Swamp

1922 . Journ. N. Y. Ento. Soc. Mar. 1922, Vol. XXX, No. i, p. 74.

1906 Dickerson, M. C. The Frog Book, pp. 131-133.

1908 Fowler, H. W. Annual Rept. N. J. S. Mus. 1906, Part I, 1907, pp. 108-112.

1908 '. Annual Rept. N. J. S. Mus. 1907, Part III, p. 193.

1930 Klots, A. B. Copeia Oct. -Dec. 1929, Jan. 16, 1930, No. 173, pp. 108-111.

1916 Miller, W. DeW. Copeia, Aug. 24, 1916, No. 34, p. 68.

1894 Moore, J. Percy. American Naturalist Dec. 1894, XXVIII, No. 336, pp. 1045, 1046.

1923 Noble, G. K. and R. C. Noble, The Anderson Tree Frog. Zoologica, Aug. 20, 1923.
1889 Peters, John E. American Naturalist Jan. 1889, Vol. XXIII. No. 265, pp. 58, 59.

Smith, W. H.
1923 Stejneger, L. and Barbour, T. A Check List of North American Amphibians and

Reptiles. Second Edition, Cambridge 1923, p. 29.
1901 Stone, W. Proc. Acad. Nat. Sci. Phila., June 1901, Vol. 53, p. 342.
1906 . American Naturalist Mar. 1906, Vol. XL, No. 471, p. 163.

Hyla cinerea (Schneider)

(PL II, Fig. 5; V, Fig. I ; VII, Figs, i, 4; X, Fig. 2; XII, Fig. 4; XV, Fig. 2; XVII; XXVI;

Text Fig. I, 12)

COMMON NAMES

Green Tree Frog. Carolina Tree Frog. Cinereous Frog. Bell Frog. Fried
Bacon Frog. Cow-bell Frog. Bull Frog. Tree Frog (Holbrook). Green Tree
Toad. Carolina Hyla. Banded Hyla {semifasciata). Hallowell's Tree Frog.

RANGE

Check list. Type Locality: "Inhabits Carolina." Range: Virginia to
Florida, west to Texas, and northward up the Mississippi Valley to southern
Illinois." Stejneger & Barbour Check List 1923, p. 30.

Supplementary records. In 1926 (Wright '26, pp. 82, 83) held Hyla
cinerea to be a Lower Austral species from southern New Jersey to Texas.
In 191 7 our route from Washington southward to Texas was through Upper
Austral or Piedmont country. We did not see or hear this species until
Flat wood (Wilco Co.) Alabama, was reached. We next took them at Pass
Christian, Miss., June 14, at Devers and Berwick, Louisiana, June 18 and
19, between Beaumont and Neches River, Texas, June 21, or from Alabama
to Texas. In Texas in 1925 we found them south of Beeville and almost to
Corpus Christi. Pope (1919, pp. 95, 96) recorded it at Houston, Tex. In
North Carolina Brmley records it in Dare and Lenoir Counties. In 1922
we found it as far west as Richmond County south of Rockingham, N. C.
In 1924 George S. Myers (1924, p. 60) reports it from Wilmington, N. C.
The following month, 1924, Schmidt reports it from Charleston, S. C. In
1922 from near Columbia, S. C, June 9, through Millen, Ga., June 10, and
Screven, Ga., June 11, and Waycross, June 12, to Okefinokee, June 13, we
recorded this species. All in all, the records distinctly make it a Lower
Austral form.

Local Okefinokee records. We recorded it from the following Okefinokee
localities: Billy's, Black Jack, Chesser, Floyd's, Honey Islands, the Pocket,
Craven's and Mixon Hammocks, Billy's and Minne Lakes, Chase and Grand
Prairies, Suwannee Canal, Suwannee and St. Mary's River, Starling Branch
and numerous localities around the swamp. It was widespread in this region
and a very common frog.

Hyla cinerea 249

GENERAL APPEARANCE

DeKay (1842 Part III p. 72) held this extra Hmital frog to be "Bright
green, with a yellow line on each side from the snout to the posterior extrem-
ities. Length if inches. From Lat. 30° N. to Mississippi."

In 1906 Dickerson (p. 127) wrote "The Green Tree Frog is perhaps the
most beautiful tree frog of North America. Its slender form and smooth fine
coat of green and gold certainly give it an air of distinction. It is more slender
than any other North American tree frog. The slenderness is accentuated
also by the lengthwise stripes of metallic white or yellow."

In 191 5 Deckert (p. 3) described it as "an aristocratic looking tree toad,
with its long, slender figure of the brightest green, edged on each side with a
band of pale gold or silvery white. Its size is larger than any of the preceding
species (//. pickenngii, H. squirella and H. femoralis) often reaching 2^
inches from snout to vent."

It is a very slim, smooth, green frog. It may have side stripe, pointed head
and shallow face ( H. cinerea) or have no stripe, broader head and deeper face
(H. c. evittata). It is relatively the longest legged Hyla of the swamp. Speci-
mens of Hyla versicolor, Hyla gratiosa and Hyla cinerea at 56 mm. have
hindlimbs respectively 70 mm., 73 mm., and 89 mm. in length.

COLORATION OF SPIRIT SPECIMENS (1912)

Color in spirits olive brown, olive green, brownish; in life all of the speci-
mens were grass green or yellowish green. The same color occurs on the dorsal
surfaces of the fore- and hindlimbs and forward from the arm on to either
side of the throat. Sometimes this narrow area of green is absent but usually
it reaches along either ramus as a narrow band seldom reaching the symphysis
of the lower jaw. In three examples (Nos. 6524, 6535, 6532), the green
extends across the throat while in the others the throat is clear. The char-
acteristic straw or yellowish band on the sides of the body is more or less
variable. In most it starts on the snout but few have it beginning beneath
the eye as Holbrook records. Usually where the tympanic fold crosses it the
band may be narrower, broken or missing for a short distance. In the ma-
jority the band reaches to the groin although several have it ending midway
between the fore- and hind-limbs. The band usually reaches its greatest
width on the side of the body. Often it is a narrow line on the sides, in one
(No. 651 1) it is almost absent and very thin being only .5 mm. long beneath
the tympanum and .5 mm. long back of the tympanic fold. This is a very
close approach to the condition of Hyla evittata.

A similarly covered line occurs along the posterior edge of the forearm
and carpus, along the posterior margin of the tibia, and tarsus and anterior
margin of the tibia ; rarely one or all of these bands may give way to green body
colors, the anterior tibial band being most frequently lost and the tarsal
carpal and forearm bands are seldom lost.

Most of the 23 adults have few to 25 small irregularly arranged cream or
orange yellow spots on the green back. In a few they become quite con-

250 Frogs of the Okefinokee Swamp

spicuous; others have the back absolutely free of them. The short tran verse
light line above the anus usually but not always present. The ventral sur-
face of the body is white or yellowish white.

STRUCTURAL CHARACTERS (WRITTEN IN 191 2)

Upper surfaces quite smooth or minutely granulated; slightly granulate
on the throat; most of the gular sac, the breast and its fold, under sides of the
arm and legs smooth; belly and posterior ventral surface of the thigh strongly
granulate. Third finger longest ; fourth toe longest and with two phalanges
free; a tympanic fold from tympanum to base of arm; vomerine teeth be-
tween inner choanae at equal intervals from them and from each other; in
two not transverse but oblique in position and in one (No. 6530) a continuous
ridge not divided into two; tongue roundish or elliptical shghtly notched;
nostrils decidedly nearer tip of snout than eye.

MEASUREMENTS

(Recent Material)

//"earf to angle of mouth i.o (20 mm.) — 1.25 (28 mm.) — 1.05 (36mm.c?') —
1. 18 (44 mm.cf ) — 1.2 (44 mm. 9 ) — 1.05 (56 mm.cf ) — i.ii (56 mm. 9 ) in
width of head; head to rear of tympanum 1.0-1.0-.95-1.0-.92-1.03-1.0-1.0
in width of head; head to angle of mouth 2.85-3.5-3.8-4.0-3.66-3.3-3.3 in
length of body; head to rear of tympanum 2.85-2.8-3.4-3. 14-3. 14-3. 1-3.0 in
length of body; snout .66-.70-.8-.92-.86-.72- 65 in first finger; snout .83-
i.o-i. 2-1. 23-1. 06-1. 05-1.0 in fourth finger; snout .66-70-1. 0-.77-.66-.77-.8
in first toe; eye i.o-i. 66-1. 25-1.3-1. 875-1. 5-1. 66 in snout; eye .5-83-75-
.50-.625-.66-.75 in tympanum; eye .66-1. 16-1.0-1. 2-1. 61-1. 08-1. 08 in first
finger; tympanum 4.0-4.2-3.0-4.4-4.8-3.75-3.33 in intertympanic width;
tympanum 2.0-2.0-1.66-2.4-3.0-2.25-2.2 in snout; internasal width i. 0-1.5-
.83-.75-1.0-.90-1.1 in upper eyelid width; interorbital width .60-.85-.71-.75-
.77-.81-.71 in upper eyelid width; interorbital width .60-.57-.8-1.0-.77-.91-
.50 in internasal width; interorbital width 2.3-2.4-2.5-2.75-2.6-2.7-2.14 in
intertympanic width.

Forelimb: Forelimb 2. i-i. 8-2. 05-1. 9-1. 83-2. 43-2. 24 in length of body;
forelimb 3.0-3.03-2.8-2.74-2.62-3.6-3.2 in hind limb; first finger i. 75-1. 57-
i-7S-i-5-i-38-1.7-1.77 in third finger; second finger i. 4-1. 22-1. 4-1. 28-1. 2-
1. 3-1. 27 in third finger; second finger .8-.77-.8-.875-.86-.76-.70 on first
finger; third finger .85-1.09-.92-.83-.88-1.0-.95 in second toe; fourth finger
1.4-1.1-1.16-1.12-1.12-1.15-1.15 in third finger; fourth finger .8-.7-.66-.75-
.81-.7-.65 in first finger; internasal width i. 33-1. 75-1. 33-1. 5-1. 85-1. 3-1. 4 in
first finger; internasal width 1.66-2. 2 5-1. 66-1. 7 5-2. i-i. 7-2.0 in second finger;
internasal width 2.33-2.75-2.33-2.25-2.5-2.2-2.5 in third finger; internasal
width 1. 66-2. 6-2.0-2. 0-2. 3-1. 9-2. 2 in fourth finger.

Hindlimh: length i. 42-1. 76-1. 41-1. 43-1. 43-1. 5-1. 43 in hind limb; tibia
2.0-1. 64-1. 94-1. 91-1. 76-1. 93-1. 86 in length; tibia 2.85-2.76-1.94-2.73-2.52-
2.9-2.66 in hind limb; tibia .95-.9-.94-1.0-.96-.80-.83 in forelimb; tibia .7-

Hyla cinerea 251

.7-.62-.78-.72-.8-. 70 in hind foot; first toe i. 5-1. 71-1. 3-1. 5-1. 6-1. 57-1. 37 in
second toe; first toe 3.0-2.4-1 .8-2.4-2.5-2. 14-2.0 in third toe; first toe 3.75-
2.85-2.4-3.2-2.85-2.37 in fourth toe; first toe 2. 75-2. 28-1. 7-2. 2-2. 2-2. 0-1.875
in fifth toe; second toe 2.0-1.41-1.38-1.6-1.56-1.36-1.45 in third toe; second
toe 2. 5-1. 66-1. 84-2. 13-2. o-i. 8-1. 72 in fourth toe; second toe i. 85-2. 28-1. 7-
1. 6-1. 375-1. 27-1.45 in fifth toe; third toe i. 25-1. 17-1. 33-1. 33-1. 28-1.33-
1.2 in fourth toe; third toe .91-.94-.94-.91-.88-.93-.93 in fifth toe; fourth
toe .93-1.15-1.12-1.12-1.12-1.15-1.1 in hind foot; fourth toe i. 33-1. 7-1. 54-
1.43-1.56-1.45-1.57 in tibia; fourth toe i. 3-1. 55-1. 46-1.43-1. 5-1. 15-1.3 in
forehmb; fifth toe i. 36-1. 25-1. 41-1. 45-1. 45-1. 42-1. 26 in fourth toe; inter-
nasal width 1. 33-1. 75-1. 66-1. 25-1. 4-1. 4-1. 77 in first toe; internasal width
2.0-3.0-2. 16-1. 875-2. 28-2. 2-2. 4 in second toe; internasal width 4.0-4.25-
3.0-3.0-3.5-3.0-3.55 in third toe; internasal width 6.0-5.0-4.0-4.0-4.5-4.0-
4.25 in fourth toe; internasal width 3.66-4.0-2.83-2.75-3.1-2.8-3.4 in fifth toe.

HABITAT

LeConte (1825, pp. 279) gives its habitat as follows: "Inhabits in great
numbers in Carolina and Georgia, particularly on water plants, such as
Pontederia cordata, the Nymphaea, and others, that generally grow in ditches,
and on the margins of rice fields." Holbrook (1842, Vol. IV, p. 120) records
that "These animals are found on trees, but most commonly about the broad-
leaved plants, as the Pontederia cordata, Nymphaea, etc; they are also very
numerous in fields of Indian corn {Zea mats), where they conceal themselves
from their enemies by passing in between the broad green leaves of the plant,
the colour of which is so nearly their own, that it is not easy to find them."
In 1856 LeConte (p. 428) records that "In the spring it is found in rice fields
in astonishing quantities."

Deckert (1914, No. 3, p. 3) "found (them) in scattered bands, among
water hyacinths on creeks." In 19 15 Deckert (p. 4) "occasionally met with
(it) on corn fields, its chief haunts, however, seem to be the shores of the
creeks, where it occurs in scattered companies on the water hyacinths and
bulrushes."

In Louisiana Viosca (1923, p. 39) finds that the denser areas in the fresh
water marshes adjacent to the wooded swamps "furnish the ideal habitat for
Hyla cinerea and Rano sphenocephala. ..."

In 1 9 1 2 we made the following notes about their habitat in the Okefinokee
Swamp: This tree frog is very common in the Okefinokee Swamp. We found
them in the large bay and cypress trees along the watercourses and about
Billy's Island. In some of the swampy edges of the islands and about Lee's
clearing they also occurred. On Honey Island Prairie the trees seemed to
teem with them during the evening chorus. The following day in the fore-
noon when the sun was very hot they were frequently taken and more often
seen on the lily-pads and other aquatic plants at the surface of the prairie.
Some were secured on the bushes and trees near our camp, on low heaths and
rarely near or on the ground.

252

Frogs of the Okefinokee Swamp

In 192 1 we found them commonly in cypress ponds, cypress bays, in
hammocks near the cypress bays or near the edge of the island, along the edge
of water courses and on the prairies. On June 4 we found a pair in the "lumber
camp quarters near a house corner on the south end of the easternmost street
in a temporary pool. Probably from houses near by. Boys tell about finding
them in or around the rain barrels." Our first specimen in 192 1, April 22, we
found in a rain barrel.

In several instances we have records of them from ivy covered porches
where they reside some times. On July 22, 192 1, at Hampton Mizell's
residence, Hopkins, Georgia, on his porch were two Hijla cinerea, one of which
we saw calling.

In 1922 we heard them quite commonly on the prairies, and in some cypress
ponds. We occasionally heard them in branches (like Starling Branch) and
in the water pits beneath the oaks and trees at Camp Pinckney, St. Mary's
River. At night the frogs may be on the bushes and stems above the water or
more frequently at the water's level. During the day they rest mainly on the
stems above the water. For example, at Camp Pinckney in the afternoon of
June 27, 1922, "took Hyla cinerea in Pond No. 2 on a vertical wild rice stem,
the frog some 3 feet above the water." Eight days later, July 5, at Camp
Pinckney, 'over Pond No. 3 on wide leaved grass-like plants were some 7 or
8 Hyla cinerea perched 1-3 feet above the water." Or two months later at the
same place August 11, 1922 "on grass stems found 3 males 2 feet above the
water."

FIRST APPEARANCE

The earliest note of activity in the South I have observed is Pope's (1919,
p. 95) record for "February 13, 19 18. Caught a young one in the grass close
to the bayou. Found it actively hopping after dark, habits evidently noc-
turnal. In the course of the next two months I collected three others, two
young and one adult 5 cm. in length. The latter was sitting on the top of a
stub close to a bayou fast asleep in the day time." In 1897 W. C. Kendall
took this species on Feb. 14, at Beecher Pt., Florida and on March 16 secured
more at the same place.

The earliest record the author has of this species is along the Medina River
south of San Antonio, Texas. Here in company with Miss Ellen D. Schultz,
Mrs. Wright, Messrs. Roy D. Quillen and R. D. Camp we found February 15,
1925, this species buried in the moist sawdust or humus or fine dust of decay-
ing logs. With it in these logs we dug out a brown snake {Potamophis striatula)
DeKay's snake {Storeria dekayi), variable swift (Sceloporus variabilis),
spiny swift (Sceloporus spinosus), ground lizard {Leiolopisma later ale), short-
lined skunk (Eumeces brevilineatus) Bufo valliceps, and TantUla nigriceps
and countless invertebrates.

GENERAL HABITS

Metachrosis. "As to coloration, this tree frog has great ability to change
from light to dark. The colour may be nearly black, or it may be so light a
greenish yellow that the stripes can scarcely be distinguished. There is some

Hyla cinerea 253

variation in colour pattern; that is, the hght bands on the legs may be nar-
rowed or obsolete, the light band on the side may extend no farther than half-
way instead of quite to the posterior end of the body, and the dark bands
bordering the light ones may be distinct or wholly lacking. . . . This Hyla
sometimes has a few small orange, black-rimmed spots on the back." (Dicker-
son, 1906, p. 127).

There is a great variation in color. In general at night yellowish green or
light greens are prevalent. During the day I have noticed pairs where the
male, the more exposed, was olive green or slaty and the less exposed female
was a hghter green than the male. Certainly the more exposed frogs of the
daytime are not always darker. The very palest Hyla cinerea I ever saw was
on the exposed edge of a cypress head, at mid-day.

The same conditions do not always produce the same coloration. In one
case we had many Hyla cinerea in a botany drum. All were light green except
three, one of which was almost black, another olive green, a dark green, and
the third yellowish green.

Variation in color. In 1825 LeConte (p. 279) wrote 'The description of
this animal should be corrected by saying, that the lateral line, from which the
name is derived, is most commonly silvery, in some few instances yellow."

In 1856 Hallowell (pp. 307, 308) held that "in lateralis {viridis Hollb.) the
lateral stripe extends as far as the anus, and there is a white band running the
whole length of the tibia, both anteriorly and posteriorly. The anterior band
is absent in semifasciataJ^ Hallowell's H. semifasciata was supposedly larger
than the true H. cmerm and had the lateral band ending half way along the side.

In 1890 and 1892, H. Garman recognized these two varieties as Hyla
cinerea cinerea and Hyla cinerea semifasciata. In 1899 G. S. Miller, Jr.,
examined the Hyla cinerea material from the vicinity of Washington and
recognized in addition to the cinerea cinerea and cinerea semifasciata a new
frog Hyla evittata "like Hyla cinerea (Daudin) but with broader, deeper
muzzle and normally unstriped body and legs." "As to the constancy of the
color differences between the two forms" (cinerea and evittata) he writes:

"I have handled about two dozen living and freshly killed specimens of
Hyla evittata, and have probably seen nearly as many more at a distance of
only a few feet. Among these one had a faintly developed stripe at the angle
of the jaw. Of the twenty-two alcoholic specimens collected by Mr. Hay and
now in the National Museum, eight have traces of the body stripe, which,
however, in no instance is margined with black, or as sharply defined as those
southern specimens in which the stripe is shortened and narrowed. Of sixty-
one specimens of Hyla cinerea (seven received alive from H. H. and C. S.
Brimley taken at Bay St. Louis, Miss., and the others preserved in alcohol
in the U. S. National Museum) there is considerable variation in the leg
stripes, though varying in length and breadth, is conspicuously developed,
definite in outline, and usually margined with black. In the two abnormal
individuals (one from Bay St. Louis, Miss., the other from New Orleans,
La.) the leg stripes are absent, and the body stripes reduced to mere traces
near the angle of the jaw. When forwarding the unstriped specimen from

2 54 Frogs of the Okefinokee Swamp

Mississippi, the Messrs. Brimley wrote that it was the only one of the kind
observed among the large numbers that have passed through their hands.
Such individuals as these are readily distinguished from the faintly striped
specimens of Hyla evittata by the form of the muzzle."

In 1 910 C. S. Brimley writes that "A single specimen resembling cinerea
but lacking the yellow line on the side, also came from here" (Bay St. Louis,
Miss.). He probably alludes to Miller's specimen. In 1915, Gronberger
writes of H. evittata as follows: "Efforts will shortly be made to secure speci-
mens of Hyla evittata Miller which are said to be very numerous in the ponds
between Mr. Palmer's farm and the Potomac River, and also to establish
whether Hyla cinerea coexists with or is not, as some claim specifically identical
with Hyla evittata.^'

In 1924 (p. 60), G. S. Myers discusses some of his material from the Wil-
mington (N. C.) country thus: "In the 14 adults, six showed partial or total
lack of the yellow band along the jaw, side and tibia. Two were altogether
without these markings, appearing very Hke H. evittata Miller. The head
shape was typical of cinerea, as observed in other specimens from Florida and
Louisiana. In the opinion of some, the blunt head of evittata, described and
figured by Miss Dickerson (Frog Book, pp. 128-130, PI. LII) is not a good
character, and evittata is simply the unhanded form of cinerea.

"With this view I cannot at present agree. Numerous times several of the
cinerea were in full cry in the terrarium and their call was very different from
that described by Miller for evittata. ..."

In 1910 Brimley (p. 11) speaks of a "Hyla (sp.) Bay St. Louis, Mississippi,
in January and February, 1901, several specimens of a tree frog with the
sharply defined side line of cinerea but with dark spots on the back like
squirella were received from this place. My notes state that they were ap-
parently a little stouter than cinerea and grayer in color."

On June 14, 191 7, several of us (Professors H. H. Knight, P. A. Munz,
G. B. Upton and others) encountered this same mixture of characters. At
the time the writer had forgotten Brimley's experience but it immediately
came to mind when the variations began to appear. The rough journal note
runs thus: "At noon we stopped in a low woods near one of the bays between
Pass Christian and Bay St. Louis. On the saw palmetto leaves we found no
end of Hyla cinerea and Hyla squirella in all color phases. Believe Hyla
andersonii not so different. These creatures were on the saw palmetto leaves
or in the bases between two stems. In one place found two specimens, one a
brown phase and one a green phase. Some have a yellowish or orange color
on the posterior faces of thighs. Others have purplish. Some have a yellowish
line on lip and a few besides have faint yellowish line on side. Many have a
purplish area from nostril to femur along the side like Hijla andersonii only
not so distinct. Adult H. cinerea were often on the face of the leaf and on
upper side. Others were on the underside of the leaf. Sometimes in one
clump of 4 or 5 plants as many as 6 or 8 specimens were found. Often a fine
correspondence of brown specimens and brown leaves or green frogs with
green blades. In one clump one yellowish green on back, another dark greenish

Plate XXVI

Green tree-frog ( Hyla cinerea)

1 . Bonnet-filled bayou on north side of Billy Lake, Ga. May 4, 1921.

2. Male croaking, Billy Id., Ga. May 24, 1921. Flashlight, x 0.35.

3. Male croaking, Billy Id., Ga. June 24, 1921. Flashlight, x 0.35.

4. Male (dark) and female, Billy Id., Ga. May 18, 192 1. Lateral aspect, x 0.33.

5. Male croaking on lily pad, Billy Id., Ga. May 16, 1921. Flashlight, x 0.33.

6. Male and female, Billy Id., Ga. April 26, 192 1. Ventral aspect, x 0.33.

7. Small tadpole, Billy Id., Ga. August 4, 1922.

8. Mature tadpole, Chesser Id., Ga. August 12, 1922. x i.o.

Hyla cinerea 255

and a third purplish brown. One captured by H. H. Knight had quite a
purpHsh area from nostril to femur. Some of the Hyla cinerea have a narrow
purplish line above and below the white, cream colored, or straw colored
lateral band. Is this the cradle of Southeastern tree frogs? There were un-
spotted H. evittata, Hyla squirella, H. cinerea, olive or green or possibly
brownish with or without lateral bands, with or without dark borders to band,
also specimens very suggestive of how Hyla andersonii form might possibly be
derived." Alas this most interesting series, one the of largest we ever collected,
dried up and was lost before we ever studied it critically. Only 14 clear Hyla
cinerea of this series remain. The interesting .specimens are lost.

In 1922 on our way to Okefinokee Swamp we heard from our camp on the
Alexandria (Henshaw Sparrow station) heights what the writer took to be
Hyla cinerea. I suspected they were Hyla evittata and Mr. Harper recognized
them as such from his previous collection of them. They impressed me as
Hyla cinerea in call and the author has always inclined to the belief that Hyla
evittata is Hyla cinerea but it is not yet settled. To call it Hyla cinerea evittata
as Dunn (19 18, p. 21) and Stejneger and Barbour (1923, p. 30) have is about
all that we can do until the life history is worked out by some one who knows
Hyla cinerea intimately. These forms from Virginia northward tend to
predominate in evittata coloration; those from North Carolina southward to
Texas are regularly striped with occasional evittata forms.

The lateral stripe present or absent is not in the southern forms a sex char-
acter. In our 192 1 material three of our records are of females 57 mm.,
55 mm., and 56 mm. in length and two males 58 mm. and 55 mm. All these
five have a jaw stripe. In a male and a female the lateral stripe is missing on
each side. In one specimen, male, the stripe is present on one side and missing
on the other side. In one female lateral stripe missing on one side. On this
side jaw stripe only 7 mm. long from beneath eye to angle of mouth. On the
opposite side the lateral stripe is just a httle beyond shoulder. In the
other female no stripe on one side and hardly none of the other side.

The head characters are supposed to be diagnostic but broader noses and
deeper faces occur with the narrow noses, nearer nostrils and shallow faces.
We have before us an evittata female with deep face, wider separated nostrils,
broader, blunter nose and a male of the same length vittate, pointed nose,
shallow face and nostrils nearer together. At transformation the stripe may
sometimes be absent; in fact at times it may be more absent than present.

General habits. In 1842 (Vol. IV, p. 121) Holbrook remarked that "At
certain seasons they may be seen on the ground on their way to pools of water;
they are exceedingly active, leaping at times eight to ten feet."

"The Green Tree Frog is most interesting when kept in captivity. It lives
high among the ferns and vines of its moss-garden, and is especially fond of
flies. It will see a fly at a distance of three or four feet, and will catch it,
making but one leap over the intervening distance. It is said that this tree
frog can leap a distance of eight to ten feet. When the call is given, the throat
pouch is inflated and the body over the lungs swells and relaxes forcibly."
(Dickerson 1906, p. 127).

256 Frogs of the Okefinokee Swamp

In 19 1 5 Deckert (p. 4) held that "Although (it was) a powerful leaper, it
is easily caught when once located, as it is not at all shy." Pope (1919, p. 96)
held "from the frequency of its call this Hyla seems to be fairly abundant
(at Houston, Tex.), but its secretive habits and protective coloration make it
very hard to find."

VOICE

In Daudin (1803, p. 29) we find that this species "croaks without ceasing
tchit-tchit-tchit-tchit." J. LeConte (1825, p. 279) records that "They are
very noisy, particularly at night, their voice somewhat resembles the sound of
a bell." LeConte (1856, p. 428) said it "is commonly known as the bell-frog,
its notes resembling the sound of small bells. It is frequently called fried
bacon frog, as its voice seems to some ears to repeat these words." Holbrook
(1842, Vol. IV, pp. 120, 121) holds "they remain quiet and motionless during
the heat of the day, but in the morning or evening, or before a shower, they
emerge from their hiding places and become very brisk and noisy. Their
noise proceeds from a single note, which, at a little distance, is not unlike the
sound of a small bell ; and there seems in general to be one leader of their or-
chestra, and when he raises his note, hundreds take it up from all parts of the
cornfield, and when he stops, the concert is at an end, until he again begins."

Harrison Carman (1892, pp. 347, 348) holds "Its note resembles the tone
of a cow-bell heard at a distance. Where abundant about water, the frogs are
very noisy just after dusk, the chorus being broken, however, by longer and
shorter intervals of silence. A single note is first heard, and as if that were a
signal, it is taken up and repeated by a dozen noisy throats till the air is
resonant with sound. After a time it ceases as suddenly as it began, to be
again resumed after a period of quiet."

In 191 5 Deckert (p. 4) says "Its call sounds like: Grab, grab, grabit,
grabit, etc., uttered in a shrill, loud voice, and has a startling resemblance to
the human voice. One would never think the call came from a frog."

In 1919 Pope (pp. 95, 95) "throughout the month of May and into June. . .
heard this frog quite often, calling from the trees at all times of day, but most
frequently late in the afternoon. The call has been compared to a cow-bell,
which it resembles in tone, but the manner of delivery suggests a cuckoo.
It usually consists of fifteen or twenty notes given in rather rapid succession,
but too distinctly to be called a trill. The cadence becomes slower after the
first, and the last few notes are often given with some hesitation. From the
frequency of the calls this Hyla seems to be fairly abundant. ..."

In 19 1 2 when we first made the acquaintance of this form in the Okefinokee
we wrote "In the daytime when the weather is sultry or especially in the eve-
nings of late May or early June some of their immense choruses are not easily
to be forgotten. Sometimes a chorus starts suddenly, quickly reaches its
crest and ends abruptly to be resumed later after a shorter or longer sharply
defined interval. Along some of the water-courses like Billy's Lake one lone
frog nearby will begin, then stop, but ere he has finished, another just ahead
of the speeding boat has taken up the task. Thus, the chorus may travel

Hyla cinerea 257

along the margin of a lake for considerable distances. Usually these experi-
ences came in the later afternoon (after 3 :oo p. m.)"

Five years elapsed before we heard them again. At Flatwood, Ala., after
6:30 p. m. ''in a drying-up swampy pond heard a chorus which sounded Hke a
cow-bell at a distance. To one member of the party it sounded like an ex-
haust running into an oil well pipe. Took one with flashlight. It was Hyla
cinerea. It was on some bushes and its sides and throat looked like a pink ball."

In 1921 we made the following notes: On Billy's Lake at night May 11,
"the Hyla cinerea of a place would sing, then stop, and the wave of sound
pass on down the lake. The frogs were mainly in the bushes along the edges
and hard to find in a casual way. We had to get out of the boat and search.
We found one two feet from water on a horizontal branch. It was literally
covered with gnats, mosquitoes, etc. After much manoeuvering, we finally
had it focused. After we had waited some time for it to croak, we discovered
the frog gone. About 11:00 p. m. the Hyla cinerea stopped. Temperature
71°. Falling temperature and cooling rain anticipated."

On May 14 at 8:00 p. m. went to the C. A. G. (Hyla cinerea, Acris gryllus,
Rana grylio) pond. When I approached the pond heard no end of Hyla
cinerea caHing everywhere. On one iris blade found a male one foot above
the water. Nearby on another iris blade was a second male. In amongst a
mat of pickerel weed (Pontederia) and lizard's-tail (Saururus) found a male
at the surface and croaking; in a bush three feet above water another male;
in a cypress tree four feet above water another; on a lily pad still another.
No females seen. When one catches them with throat inflated they give a
funny squeak as if the balloon had burst.

When the chorus is in full swing the males are easy to capture, not hard
to observe or photograph by flashlight. On April 25, 192 1, we went to one
pond "where many Hyla cinerea were on lily pads. Several were also in the
bushes. One of these allowed me to stroke it with a flashlight and did not
move. Finally it merely scrambled farther down the branch on which it was
resting." At other times it will leap from the branch to another lower branch
and will land in the water. In the latter case it scurries along on the water to
some vegetation for a new perch.

This is easily one of the most characteristic anuran voices of the swamp.
We have heard it from mid April to September. Its call is loud, the frogs are
numerous and the voice period is long. It is one of the rain signals for the
residents who call it "rain frog."

Mr. F. Harper makes the following notes on its voice: "The calls are given
at the rate of about 75 per minute: quonk, quonk, quonk, quonk, etc. Mean-
while there is an alternate inflation and deflation of throat and body. The
note comes at the maximum inflation of the throat sac, which is then fully
half the size of the body. Between croaks the throat does not fully collapse,
there remaining a little balloon-like expansion in the middle. After starting
off with the ordinary note described above, a frog will often redouble its
speed and at the same time add a rolling quality to the note: crronk-crronk-
crronk-crronk, etc. The throat sac is then only slightly deflated between

2S8

Frogs of the Okefinokee Swamp

croaks. When two individuals are calling together, but not in unison, there
is an interesting antiphonal effect, as of a double note: bo-habe, bo-babe, bo-
babe, etc. Probably both the calling periods and the intervals between them
are irregular in length. On one occasion the former appeared to last i^ min-
utes, and the latter about 2 minutes. When one frog pipes up, after an inter-
val of general silence, all the others in the vicinity generally chime in almost
immediately, as if having just awaited such a signal to start the chorus.
As a matter of fact, a spirit of competition probably enters here, each male
being desirous of announcing his location to any nearby female that may be
ready to mate.

A resume of the croaking records for 192 1 and 1922 is:

April

April
April
April
May

May
May
May

May
May

May

25

27

30

3

9
10
II

13

14

16.

May

19

May

20

May

21

May

22

May

24

May

26

May

27

May

30

June

3

June

4

June

5

June

7

June

8

June

15

June

19

June

21

June

23

June

24.

June

25-

I92I
Heard one in Long Pond 7:30 p. m. Males on bonnets croaking in
evening in another pond.

One heard at 10:00 a. m. Few heard in afternoon.
Many heard at night.

Cold tonight. Air 58°. No Hyla cinerea calling.
Tonight at 8:00 the temperature is 56°F. The last few days have
been so cool that the Hyla cinerea beginning choruses are closed.
Billy's Lake, a few calling at 8:30 p. m. Temperature 73°.
Croaking at night the Southern bull frog pond.
"Wave" chorus at Billy's Lake stopped about 1 1 :oo p. m. Tempera-
ture fell to 71° and a cooling rain anticipated. It came.
Few heard in afternoon.

In a Pontederia and Saururus pond heard tonight no end of Hyla
cinerea.

At 7 :oo p. m. heard an immense and intense chorus along Billy's
Lake in all the ponds and bays. "We flashed a Hijla cinerea calling.
They are easy subjects."
At times plenty of Htjla cinerea calling.
Common at night.

Hyla cinerea going strong at 7 :oo p. m. Air 70°. Few calling in mid
afternoon.

Wonderful chorus east of Billy's Lake. Great racket in Moonshine
Pond 9:00 p. m.

"Strong tonight" at 10:00 p. m. Air 77°F. No end of them calling.
In many ponds the dominant note. Starting up at 5 :oo p. m.
Going 8-11:00 p. m. and later.

Out tonight. Heard H. cinerea calling commonly.
A few calling
One or two calling.

Many heard in various places at night.
Air 71°. Many heard at Moonshine Pond.
A number calling about dusk. Air 70°.

Tonight at times quite a chorus in some of the cypress ponds.
Air 8o°F.

Every night for a week Hyla cinerea has been active.
Last night a chorus.

Ponds alive with croaking H. cinerea after the evening's rain of 2
or more inches.
Hyla cinerea abundant.

Strong tonight. "Today heard several and once or twice in hot day
H. cinerea burst into chorus. Is a storm coming?"

Hyla cinerea

259

A large chorus.

Commonly heard in one pond.

A few calling from 5:00 p. m. onward.

Some yet calling at Long Pond.

A few heard.

A few early in evening and a moderate chorus later.

Some calling. Later a great chorus on prairie.

A few heard at night.

Singing at 8:00 a. m. At night calling loudly.

Large chorus.

Last night a chorus.

Many heard on prairie.

Several calling.

Hxjla cinerea calling.

Heard almost every night.

Heard in forenoon.

1922
Hyla cinerea in chorus at Everett's Pond, N. C. (near Cheraw, S.C.)
Near Millen, Ga., heard a chorus.
Heard one 6-7:30 p. m.
Great choruses on prairies.
One heard.

Plenty calling on Suwannee Canal and on prairies.
Calling loudly at Chesser Id. landing.
Heard a few last night.
Calling on prairie.

Chorus at Lake Seagrove 8:00 p. m.
Heard at Coat Bet Pond.
A few H. cinerea on prairies.

"Not so many calling on prairies as I would suppose after the hard
rain."

Heard a great breeding chorus of Hijla cinerea near Starling Branch.
One heard on Chesser Island.
One heard 9:00 p. m.
Several calling.
Several calling.

Commonly heard on Billy's Lake at night.
Several calling Chesser Island.

A very few H. cinerea calling 8-11:30 p. m. after rain. Tempera-
ture 79°.

Large chorus Camp Pinckney.

Several heard on trip to St. Mary's, Chesser Id., 7-10:30 p. m.
7 :oo p. m. H. cinerea calling. Callahan, Florida.

When the choruses of Hyla cinerea began after dusk, 7o°-82° or 84°F. tem-
perature prevailed. For example, in 192 1 this species was heard for several
days in last of April but when in early evening s8°F. was recorded on April 30
none were heard. From this date for several days in May the minima ranged
from 44°-54° until May 8. No Hyla cinerea were calhng. But on May 9 the
early evening temperatures reached to 73° and the green tree frogs began
again. Thereafter 68°-8o° temperatures generally prevailed when calling
occurred. But the great choruses came when the storms threatened, were
imminent, or after heavy downpours. Humidity is doubtless more important
than temperature in inducing calling or choruses.

June

28.

June

29.

July

3-

July

4-

July

6.

July

9-

July

II.

July

17-

July

23-

July

24.

July

26.

July

27.

July

29.

July

30.

August

1-12,

August

23-

June

8.

June

10.

June

12.

June 14

-22.

June

28.

June

29.

July

2.

July

3-

July

5-

July

7.

July

12.

July

23-

July

25-

July

26.

July

27.

July

29.

August

I.

August

2.

August

6.

August

7-

August

8.

August

II.

August

13-

August

15-

26o Frogs of the Okefinokee Swamp

MATING

Male (From, life, April 26, 1921). In dark olive green male there are twenty
orange spots scattered over the back, lighter in green of sides below lateral
stripe; stripe (cream colored) extends almost to hind legs. Obscure on snout
ahead of eye. Cream colored or white stripe on back of forearm. Iris bronzy.
Pupils horizontal.

Same on back of lower leg and along hind end of foot. Throat from angle
to mouth to slightly back of chin green. Chin proper yellowish cream. Back
of these two areas, yellowish cream and green, is the wrinkled pink area of
the throat proper.

One light colored male with stripe straw-colored, only an inch behind in-
sertion of arm area black above and below the stripe. (Non Ridgway.)

{From life, May 18, 1921). Back may be apple green, dark green olive,
greenish oUve, deep slaty olive to almost black, the stripe on side light dull
green yellow or clear dull yellow. In one male stripe not beyond tympanum
and suggestive of H. evittata. The vent, forearm stripe, foot stripe and heel
are white. One dark male had the foot stripe fainter. Iris russet vinaceous.

Female {From life, April 26, 1921). With no pink chin, green extends in
on sides of throat for slight distance. Straw colored stripe to insertion of hind
legs prominent, also stripe under eye and around stripe. Under parts from
groin to chin the same cream color. (Non Ridgway).

{From life, May 18, 1921.) Back may be from apple green to deep slaty
olive. Stripe pale chalcedony yellow.

Structural differences. In the pair of May lo, 192 1, the female was much
longer. On June 4, 192 1, the mated pair captured the day before was colored
at 11:00 a. m. as follows: Female, yellow green; male deep slaty olive. An-
other female in the jar was also deep slaty olive.

In 191 2 we analyzed the differences thus: In the series of 23 adults, with-
out exception the six females are the largest being 56, 57, 58.5, 59, 59, 60 mm.,
average 58 mm. ; while the 1 5 males range from 46-55 mm., average 5 1 mm. Of
course, the females can be told because of the absence of the vocal sac which
in the males is not especially colored. In one or two of the females the throat
was quite greenish across the whole width. The breast fold is not so promi-
nent and the throat region not wrinkled area (in spirits). A firm and full
appearance in the inguinal region reveals the ripe females at once though the
waist is almost as in the males, — slender even when the eggs are ripe. In the
male enlarged thumbs or nuptial excrescences seem to be absent.

A survey of our 192 1 and 1922 material with comparisons with our 19 12
Okefinokee specimens and other material from other parts of the range of this
species reveals two gravid females of 41 mm., two females at 44 mm. and one
at 45 mm. One would expect the male external characters to appear slightly
smaller. One 35 mm. specimen has a slightly wrinkled throat. One specimen
of 37 mm. has the throat wrinkled and is a male. Of three 38 mm. specimens
one is doubtfully male externally; another has the throat wrinkled but not
prominently and is a distinct male. Two specimens 39 and 42 mm. appear to
be males. From 44 mm. onward we have many males. The evidence if sum-

Hyla cinerea 261

marized is: 28 females from 41-63 mm. in length, average 53 mm., mode 57;
53 males from 37-59 mm. in length, average 50 mm., mode 55 mm. The
males and females may begin breeding rarely at the end of their second year
but the rate of growth and ripening of gonads point to the beginning of the
third year and third spring after transformation as the first ovulation for the
individual frogs.

Duration, day or night. The first record we have is on the evening of May
10, 1921. "This evening at 8:00 p. m. (Temperature 72°) went to new pond
southwest of boys' swimming hole in Billy's Lake. Here were no end oi Acris
gryllus and Rana grylio in all stages. Easy to capture them at night except
the adult R. grylio but even these can be taken. The pond is cypress-lined.
Moon shows just above them. Beautiful picture though some might think of
the moccasins, alligators, etc. when wading around at night. Heard a Hyla
cinerea nearby. Began to search. It stopped. Later thought I saw it.
Seized it and it proved a ripe female, not croaker. This female was in a
cypress 4 feet above the water. Went away. Later the croaker started
again. On the flat side of an iris leaf three feet from the captured female was
the male. Was the female approaching the croaker? Put both in a jar.
Came back to camp. The male and female are mated at 9:00 p. m." On the
morning of May 1 1 the pair were broken and no eggs laid.

On the night of July 3, 192 1, after a heavy rain we found two mated pairs
of Hyla cinerea not in their accustomed place of ovulation. In the water
between the furrows of sweet potatoes found one of the pairs. The other
was in another cultivated field between furrows of cane or corn. Biifo quer-
cicus and Hyla jemoralis pairs were also in numbers.

On July 7, another pair was found in a temporary open grassy pool near
the lumber company's wood pile, quite an aberrant place for ovulation.
Equally so was the temporary pool in the street of the Billy Island Lumber
Camp where on June 4, 192 1, a pair was found.

As to duration of embrace we have httle evidence. Normally they lay the
same night a pair is captured. The pair of May 10 were broken the morning
of May II. On the night of May 14 these were mated again and then alter-
nately broke and embraced for several days after without ovulation.

Amplexation (Normal, abnormal). The pair mated May 10, 192 1, had
an axillary embrace. "The bent toe pads of two fingers (outer ones) of the
male show slightly." On May 11 they broke. On the night of May 14 "The
male held the female with his forelegs in front of the female's forelegs. One
arm came to the middle line of the female's throat and the other arm came to
the corner of her mouth. Later the male embraced normally."

On May 14, 192 1, brought in many males. Put them together and two
were soon mated in an axillary fashion. The embrace was not strong for
they broke when we tried to flashhght them.

On May 18, 1921, "at 9:30 p. m. in Long Pond on moss edge, not in pond
proper we found a mated pair of Hyla cinerea. Apparently the females are
often met at the very pond's edge. They broke. Put them in photo jar.

262 Fr'ogs of the Okefinokee Swamp

Later they resumed." The embrace was a normal axillary amplexation as it
was with the mated pair of June 4, 192 1, the two pairs of July 3, 192 1, and
the pair of July 7, 1921.

OVULATION

Habitat. Under voice we have noted this species in the bushes in the ver-
dant edges of Billy's Lake and in a cypress pond (C.A.G.) with its mats of
sphagnum, Hydrocotyle, Iris, Pontederia, Saururus, Castalia, Nymphoides,
etc. On May 19, 192 1, we made these comparative notes on habitat from
one cypress pond: "Rana clamata along outer edges in Decodon, some Pon-
tederia and entirely on east side of Long Pond. Rana grylio in middle. Acris
gryllus widespread. Hyla jemoralis mainly on either northern or southern
end. Hyla cinerea out in the middle and around edges also but Hyla jemoralis
as yet around edges." They also breed commonly on the prairies, sometimes
in the cypress bays and at times in transient pools or inundated cultivated
fields of the islands or mainland.

Period. In 1915, at Jacksonville, Florida, Deckert (p. 4) reported that
"This species seems to have no special breeding season, and is never heard in
the early spring, when the majority of the other frogs and toads congregate
at the shallow bayous." Pope (1919, p. 96) of Houston, wrote 'T could get
no light on its breeding habits."

From the evidence of mated pairs in 192 1 we might call May 10- July 7
the maximum period. In 192 1 we recorded eggs May 18, May 23, June 4,
July 3 or from May 18- July 3.

In 1923 Wright (1923, p. 34) held that "From May 15th to June ist five
more (species) start (ovulation). . . . Hylacinerea . . ." In 1924 Wright and
Wright (1924, p. 378) give the "Season, May 19 to August 21" as the period
of ovulation. Another note represents breeding as beginning May 10 and still
another notation has the period from May 18- July 30.

In 19 1 2 we evaluated our evidence as follows: "We suspected this species
must be breeding during the period of the greatest choruses which we re-
corded May 30- June 4. Of the 12 adults taken between May 30- July 15,
19 1 2, only two were females (taken respectively June 3 and June 6). Only
the first of these two was gravid, the other was spent. Of the ten adults
secured after July 15, four proved ripe females. Then, the remains of an-
other frog taken June 15, 19 12, from the stomach of a black-snake proved
to be a ripe female. Thus we have outhned a period of ripe females from
June 3- July 1 5 or later. The species may breed from the last of May through
June and July."

A summary of our material on the basis of spent and unspent females for
years not of 19 12 is as follows: On April 28, 192 1, the only two females caught
respectively 55 mm. and 57 mm. are spent. On May 15, 192 1, we took three
females, two 55 mm. and 56 mm. spent, and one 57 mm., gravid. In June,
191 1, at Spring Creek, Decatur Co., Ga., Professor J. C. Bradley took two
females 44 and 59 mm. respectively, each gravid. On June 5, 192 1, we se-
cured a ripe female 59 mm. long. The day before another unspent female 57

Hyla cinerea 263

mm. On June 21, 19 17, at Devers, Texas, we captured a gravid female.
On June 30, 192 1, a female 56 mm. was not quite ripe. On July 3, 192 1, we
have two spent females.

This evidence of 191 1, 19 17, and 192 1 yields the facts that breeding began
before April 28 and reached at least to July 3. The gravid females recorded
above for the period after July 15, 19 12, send the ovulation far into July.
From spent and unspent females evidence, mating pairs, actual eggs recorded
and early and late choruses this species may lay at least from mid April to
mid August or later.

Temperature and humidity. In the six or seven instances of ovulation or
mating in 1921, we have a rain of .88-2.80 inches before or .84-1.00 the day
of the record. On the day before the record the maxima ranged from 75°-9i°,
the minima 60^-74°; on the day of the record, maxima 8o°-9i°; minima 62°-
70°. Close study of these temperatures shows 65°-76° minima prevalent
when ovulation occurred.

On May 16, 192 1, when we had an immense chorus we made the note
"The humidity, previous rain and threatening rain must be more of a factor
than temperature."

Egg-laying process. In 19 12 if the presumptive egg packets (June i,
191 2) of 5-30 eggs were H. cinerea these must be moving about as in Hyla
versicolor, Hyla femoralis and some others. Then we held that each female
if she lays packets of only five each would have about 70-100 packets to lay
or if in masses of 30 each about 10-16 different ovipositions. In 192 1 on May
23 when we found them in the field sometimes the packets seemed distinct
and other times not so much so. The eggs might be more or less strewn on
the vegetation or at other times be in packets. Then we noted on the spot
that all the complement apparently is not laid at the same place or time; there
is some moving about. Possibly its method may be somewhat akin to that
described by Noble and Noble for Hyla andersonii where it strews its eggs on
the sphagnum.

The eggs laid May 18, 192 1, were not hatched May 24, 192 1. They
eventually spoiled. Doubtless this means they were laid by the female when
not mated. On July 9, 192 1, we also recorded that the eggs laid by the pair of
July 4 did not hatch. Here also they were doubtless laid when not paired.
If each case represents infertility from male it may be the captivity, restricted
quarters where there is little opportunity to move about.

EGGS

Attachment, egg mass. The first eggs laid by the pair of May 18, 192 1,
were deposited in our photographic jar and give no idea of the mass form in
the field. In the illustration they appear as a flocculent mass of eggs.

In 19 1 2 we made these notes on the eggs of this species: "On June i, 19 12,
Dave Lee found some amphibian spawn in some pools of their clearing.
These pools were near the swampy thicket around the edge of the cultivated
corn and chufa's fields and were decidedly transient in nature. The eggs
were in masses on the surface of the water more or less like tree toad {Hyla

264 Frogs of the Okefinokee Swamp

versicolor) eggs. In each mass were 5-30 eggs. These packets were easily dis-
cernible because of the excessive amount of air bubbles in them. In a few
instances the masses were slightly beneath the surface an inch or more and
the eggs matted together in semi-ball-like fashion. Usually, however, the
film form of the tree toad complement obtains with this species. In addition
the color of the eggs (brown animal pole and cream or yellowish white vege-
tative pole) makes the resemblance more marked. A comparison of the eggs
taken in the field with the ovarian eggs of Hyla carolinensis satisfies the
authors that the identification is correct but it is yet circumstantial or pre-
sumptive." Now we know the presumptive evidence would be just as strong
or stronger for these masses being those of H. femoralis, yet we cannot forget
that in these transient pools of these same cultivated fields we found two
mated pairs of Hyla cinerea. Doubtless they could not lay normal masses in
shallow water as they do in deeper cypress ponds, prairies or lakes.

On May 23, 1921, "from 2:30-5 p. m. we spent on Billy's Lake in a 'bon-
net' (waterlily) area where Billy's Lake turns into the run for Suwannee Canal.
Here we have heard a wonderful Hyla cinerea chorus. On the edge of a yellow
waterlily (Nuphar) leaf where a part of the edge was submerged was a mass
of H. cinerea eggs, 25 in all; in a water pocket where the petiole stem enters
the midrib of the leaf was another packet of 10 eggs. The first packet I saw
was on a bladderwort {Utricularia) stem. Another was wound around a
Nuphar petiole. In one bladderwort the eggs were amongst the leaves and
3-4 inches below the surface. Hard to distinguish the eggs and bladders at
times. Another mass attached to bladderwort leaves or floats. A little
farther on a mass amongst algae."

The later records of 1921 are not presumptive. We had the camp eggs
laid May 18-19 to compare with these field-laid eggs of May 22-23.

Egg description. Our field notes are as follows : Measurements of the eggs
laid in camp May 18-19, 1921, are: ".8 mm. vitellus; 2.8 mm. inner envelope;
3.6-4.0 mm. outer envelope. The animal pole is blackish, very sHghtly
brownish; the vegetative pole about as white as many frog's (Rana) eggs."
On June 4, 192 1, the following notes are made: "Eggs of Hyla cinerea are as
follows: vitellus i. 0-1.2 mm.; inner envelope 2.2 mm. or 2.4 mm. Animal
pole black or brownish black, darker than in Hyla femoralis, animal pole
almost white or cream color sometimes a slight yellowish tinge but not so
much as in Hyla femoralis. Outer envelope a part apparently of jelly ma-
terial and poorly defined."

Mrs. Wright's laboratory examinations give the following measurements :
vitellus .8 mm.-i.6 mm. for 29 specimens, mode 1.2 mm., average 1.15 mm.;
inner envelope 2.0-3.4 mm., mode 2.4 mm., average 2.55 mm., outer en-
velope, 3.2-5.0 mm., mode 4.0 mm., average 3.8 mm.

In 19 1 2 the count of the ripe eggs in one female gives 343 and in some of
the others where slightly more occur the complement certainly does not
exceed 500. In comparison with some of the other tree frogs it is a relatively
small complement for a frog 50-60 mm. long. Of course, the form is very
slender of body.

Hyla cinerea 265

Dangers. The presumptive eggs discovered June i, 191 2, in the edge of the
cleared fields were in a pool with 2 inches of water. On June 4, the pool was
a mere moist depression with no water. No doubt the percentage of loss
from such a cause is great but not so much so in this species as in Hyla
femoralis, Bufo quercicus, Gastrophryne carolinensis. Hyla cinerea normally-
lays in deeper water than these three species.

HATCHING PERIOD

The eggs discovered June i, 191 2, in a pool with 2 inches of water were
hatched June 4, or three days later. The eggs laid by a pair after midnight
on the night of June 4-5, 192 1, hatched June 7 by 6:00 a. m., or about two
days after ovulation. These were in a shallow basin and the air temperatures
were: maxima 82°-89°, minima 66°-7o°.

MATURE TADPOLE

Color description from life {July 26, 1921). Top of body generally citrine
or olive green or oil green, sometimes a bright spinach green or forest green.
Stripe on side of head from snout to eye sulphur yellow and ivory yellow.
Belly solid cartridge buff or ivory yellow. Breast and lower throat light
vinaceous fawn. Mental region and upper throat dusky slate-blue with scat-
tered spinach green spots. Sides of the body in branchial region and under
eye spotted dusky and slate blue or slate-blue, light vinaceous fawn, cartridge
buff, sulphur yellow.

General color of tail sulphine yellow to citrine. Amongst brown and far
down tail sulphur yellow to light green yellow edges to crests or between
crests. Base of muscular part of tail with some light vinaceous-fawn spots
along the middle of it for i centimeter or more. In almost mature tadpoles
not black spots on the crests. About time hind legs begin to grow upper and
lower crests with prominent blackish spots. Fine spinach green flecks over
tail except near the edges of the tail crests. In the younger tadpoles without
blackish spots the black is in specks all over the tail and these faintly appear
on the tail crests as dark edges.

Iris light cadmium to buffy yellow with dusky in front and behind the
pupil.

General appearance. Tadpole medium (40 mm.) full and deep bodied.
Tail acuminate, tip acuminate, sometimes acute. Tail long. The dorsal and
ventral crests about equal and about equal to musculature in depth. The
dorsal crest extending on to the body to the vertical about midway between
the eye and spiracle. Spiracle sinistral, directed upwards and backwards,
far below lateral axis, the spiracular opening very visible as a round or elHpti-
cal opening. Eye on lateral axis, in dorsal aspect on the lateral outline and
in consequence visible from the venter. Anus dextral, or only slightly above
the lower edge of the ventral crest. Muciferous crypts not distinct. As they
approach transformation the whole tail becomes spotted with conspicu-
ously dark and light spots.

2 66 Frogs of the Okefinokee Swamp

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
fringe for about 1/4 to 2/9 of the length of the upper fringe. The end of the
second row is usually even with the end of the upper fringe. The horny beak
is contained in the upper fringe 2-2.3. Median space between the lateral
second upper labial row 3 or 4 times in the length of the lateral row. The
latter row about 2 1/2 in the upper fringe. Inner papillae extend beneath the
third lower labial row of teeth often giving a two rowed appearance below it
like H. andersonii, H. femoralis, H. versicolor or H. squirella. The third
lower row of teeth, however, is not like this group but shorter like H. gratiosa
or H. crucifer. In one specimen it was absent, with the papillae below also
absent. This third row is contained 2/5-1/6 in the length of the first or second
row of lower labial teeth. The first and second rows (lower labial) are equal
or I 1/3 greater in length than horny beaks. Sometimes fringes are united
by their ends as in H. gratiosa.

Measureinents. Length of body (11-15 mm.) in tail (16-25 mm.) 1.25-2.0,
average 1.6. Width (6-7.5 mm.) of body in its own length 1.8-2.2, average
1.9, Depth of body (6.0-7.0 mm.) equals body width. Depth of body
1.69-2.5 in body length, average 1.95. Depth (7.0-8.0) of tail in length of
tail 2.0-3.125, average 2.75. Muscular part (4.0) 1.75-2.4 in depth of tail,
average 1.925. Spiracle 1. 15- 1.6 nearer base of hind legs or vent region (5.0-
7.0 mm.) than the tip of the snout (8.0-9.0 mm.), average 1.375. Spiracle
1.4-2.3 nearer eye (3.0-3.5 mm.) than base of hind legs or vent, average 1.85.
Eye 1.1-1.65 nearer spiracle (3.0-3.5 mm.) than to tip of snout (3.5-5.0 mm.),
average 1.38. Nostril i. 0-1.5 nearer eye (1.8-2.5 mm.) than snout (2.0-3.5
mm.), average 1.35. Mouth (2.0-3.5 mm.) 1.-1.4 in internasal space (3.0-4.0
mm.), average 1.25. Mouth contained 1.6-2.0 (average 1.83) in interorbital
distance (5.0-6.0 mm.). Internasal space contained in interorbital space
1. 2 5-1. 8, average 1.48.

The dimensions of the largest tadpole are :

mm.

mm.

Total length

40

Spiracle to vent

7.0

Body length

IS

Spiracle to eye

30

Body depth

6

Eye to snout

5-0

Body width

7.0

Eye to nostril

2-5

Tail length

25.0

Nostril to snout

3-5

Tail depth

8.0

Mouth

3-0

Musculature of tail

4.0

Interorbital distance

6.0

Spiracle to snout

9.0

Internasal distance

4.0

General remarks. In 1923 Wright (1923, p. 406) gives this characteriza-
tion: " Hyla cinerea — greenish tadpoles, yellowish tails." When they are
young they have a little light cross band across the body reminding one of
the pattern in small Rana heckscheri tadpoles. Before we knew this Rana's
tadpoles well we had superficially identified its young tadpoles as Hyla
cinerea tadpoles.

Hyla cinerea 267

In 1922 by living on Chesser Island we secured less of this species and
Rana grijlio but 192 1 we used to secure these two tadpoles in considerable
numbers in the floating islands of vegetation along the edge of Billy's
Island. We would haul the vegetation into the boat. Tadpoles of these two
frogs Hyla cinerea and Rana grylio were the commonest anurans. In 1922
we had great difficulty in separating young Rana grylio tadpoles from mature
Hyla cinerea tadpoles.

In 1922 the mature tadpoles were mainly taken in July and in 1922 we
recorded them July 24 and August 11. This seems to imply ovulation mainly
after the first of May.

LARVAL PERIOD

In 192 1 fully grown tadpoles were captured in July. The first eggs we
recorded were May 18, 192 1, and the first mated pair May 10, 192 1. The
first recorded transformation in 192 1 is July 12. This might imply 55 to 63
days for larval life.

TRANSFORMATION

Period. In 192 1 most of the tadpoles of July 25 implied transformation
August I and later, although six were actually at transformation. In 1922
mature tadpoles were recorded as late as July 24 and August 11. This im-
plies transformation in August or as late as September i. Surely eggs laid
July 3, 192 1, could not transform short of the last of August or September i.
Our latest date of transformation is October, 191 5. This must be from eggs
laid by August breeders. In 191 2 we recorded one July 2 from Minne Lake.
Our range of dates is from July 2 to October. July and early August doubt-
less are the crest of transformation.

Size. Our data are meager. In 19 12 one transformed frog measured 12
mm. One taken October, 191 5, was only 11.5 mm. The lone specimen of
192 1, July 12, is 15 mm. In Long Pond, Billy's Island, July 25, 192 1, we
took six transforming frogs, 16, 16, 16, 16, 16, 17 mm. respectively. One or
two tadpoles (July 25, 192 1) which were almost at transformation measured
16 mm. On July 26, 192 1, in a smaller place we found a tadpole a very few
days from transforming and 17 mm. in body length. Possibly 11.5 mm.-i7
mm. fairly outlines the transformation size.

GROWTH

In October, 1915, we recorded a 11.5 mm. transformed specimen and
October 9, 1906 a set of three 25, 27, 30 mm. respectively, two size groups.
Several isolated transformed records of 12, 15, 16, 17 mm. respectively were
made. In one set from Florida we have a 14 mm. and 20 and 26 mm. speci-
men— two groups. Another set has 23, 24, 24 mm. specimens and a 33 mm.
individual, possibly two groups. From Pass Christian, Miss., June 14, 191 7,
we have the following at 36, 38, 39, 41, 41, 42, 45, 45, 46, 46, 47, 48, 48, 48 mm.
respectively. On July 15, 19 16, we have some more specimens from the
same size group: 35, 36, 36, 37, 38, 38, 38, 39 mm. respectively. On July 14,
191 5, four specimens are at hand, one 38 mm. and three 44,45 and 47 mm. re-

268 Frogs of the Okefinokee Swamp

spectively are recorded. This evidence thus far might be interpreted in two
ways: ist years old 20-27 mm., or 20-30 mm., 2nd years old 27-39 mm. or
30-42 mm.; 3rd year olds 41-46 mm. or 44 mm. Beginning with the larger
sized series, we have the following evidence. On July 15, 1922, we took 45
mm. and 55 mm. specimens, apparently two groups, 3rd and 4th year olds.
In June, 191 1, we have 44, 44, 46, 48 mm. specimens and 55 and 59 mm.
specimens, two groups, again distinct groups. But on June 2-6, 19 12, we
have a flowing series of 47, 48, 50, 50, 52, 54, 54, 56, 58 mm. specimens. We
might interpret them all of one year 47-58 mm., or in the light of other ma-
terial break the third year olds either at 49 or 55 mm. We have nine col-
lections where none of the individuals are below 50, 51, 52 or 53 mm. and
these reach to 63. In this group 55 mm. is the mode with 57 mm. and 52 mm.
next in order of frequency. In the lower group 45 mm. and 48 mm. are the
modal points. In the 30-39 mm. ranges 38 mm. is the modal point.

If modal points be used the age groups are 11.5 mm.-i7 mm. transforma-
tion; 20-30 mm., I year olds, 24 mm. mode; 33 mm.-42 mm., 2 year olds,
38 mm. mode; 45 mm.-5o mm., 3 year olds, 48 mm. mode; 51-59 mm., 4
year olds, 55 mm. mode; 63 mm., 5 year olds. On the basis of absent meas-
urements and other data we might interpret the groups thus: of these 103
frogs II. 5-17 mm. at transformation, 17-27 mm. one year olds; 27-39 mm.
as two year olds, no 40 mm. specimen; 41-48 mm. three year olds, no 49 mm.
specimen; 50-60 mm. four year olds, no 61 or 62 mm. specimens; 63 mm.,
five year old.

FOOD

Holbrook (1842, Vol. IV, p. 121) reported that "They feed on various in-
sects, especially the common fly, which they seize with great adroitness,
leaping a foot or more to secure it."

Dr. Vernon R. Haber has examined all our material. In his paper on
"The Food of the Carolina Tree Frog, Hyla cinerea Schneider" he sum-
marizes the food as follows:

"The Carolina tree frog, Hyla cinerea Schneider, habitually is arboreal,
at least non-aquatic, excepting during mating and hibernating periods.

"The younger forms rapidly adapt themselves to a type of diet which is
characteristic of the adults of the same species, as is revealed in table 2.

"Habitually frogs of this species feed upon characteristically non-aquatic
forms until they are absolutely obHged to resort to water for breeding pur-
poses. They may take a few aquatic forms while en route to and from the
breeding places, for most of the aquatic forms could have been taken above
the water surface, as explained in the text. It is doubtful that frogs of this
species feed while mating.

"It is not hkely that these frogs while in the water for mating purposes
had taken terrestrial forms which had fallen into the water, for if such had
been the case, it seems that they also would have taken a larger percentage of
aquatic forms which habitually swim about upon or near the water surface.

"The diet of this species of frog includes such destructive forms as those
listed in table i. Of the five families of destructive Orthoptera, representa-

Hyla cinerea 269

tives of all but one, the Phasmidae or walking sticks, were recovered. Of the
68 per cent which recently had eaten insects, one-third had taken Orthoptera,
and slightly more than one-fifth had taken to Heteroptera.

92 per cent had taken arthropods.

68 per cent had eaten insects.

24 per cent had devoured Orthoptera, of which cockroaches were found in
II per cent, Acrididae in 8 per cent, Gryllidae in 3 per cent, Mantidae
and Locustidae each in i per cent.

24 per cent had but a short time previous secured Coleoptera, of which 5
per cent had eaten Cerambycidae, 4 per cent had taken CurcuHonidae,
3 per cent had devoured CisteHdae; Lampyridae, Coccinellidae and
Scarabaeidae each had been swallowed by 2 per cent, while representa-
tives of the following were included in the diet of i per cent each; Anthri-
bidae, Carabidae, ChrysomeHdae, Cicindelidae, Lathridiidae and
Staphylinidae.

21 per cent lately had gotten Lepidoptera, 7 per cent yielded caterpillar
remains.

15 per cent had devoured Hymenoptera. Ants were taken from 12 per cent
and Scoliidae from i per cent.

13 per cent had obtained true bugs, of which but 8 per cent were further
recognized. From the digestive tracts of 2 per cent each came Neptidae
and Pentatomidae, which each of the following were yielded by but i
per cent respectively; Belastomatidae, Corixidae, Pyrrhocoridae and
Scutellaridae.
7 per cent had taken Odonata.

6 per cent had eaten Diptera.

3 per cent had obtained Homoptera.

Aphidae coming from 2 per cent, Jassidae from i per cent.
2 per cent had included Physopoda or Thysanoptera.

1 per cent gave up each of the following: Euplexoptera, Neuroptera and

Trichoptera.

51 per cent a short time before they were captured had eaten harmful insects,
and several had taken several of such insects.

31 per cent recently had gotten insects which are of useful or harmful eco-
nomic importance.

15 per cent had taken insects which are of beneficial economic importance.

7 per cent recently had eaten insects which are of negligible economic value.
24 per cent positively had taken spiders, of which 6 per cent could be further

identified, 2 per cent had taken Lycosidae, 2 per cent had Salticidae,
I per cent had taken Filistatidae, and i per cent had taken Argiopidae;
this latter i per cent had eaten three argiopids all of which were easily
identified to the species. The remaining 18 per cent contained bits of
legs, chelicerae and other portions of the bodies of spiders which could
not be further identified.

4 per cent had gotten ectoparasitic mites of insects.

2 per cent had included Phalangidae.

I per cent respectively yielded each of the following: Myriapoda, Phyllopoda

and Pseudoscorpionida.
55 per cent arthropod remains which could not be further identified beyond

reasonable doubt.
36 per cent were gluttonous, see it of all tables and interpretative notes.
35 per cent had swallowed epidermis.
41 per cent were nematode infested, usually from the duodenum posteriorly;

in I per cent nematodes were found in the stomach. Most of the nema-

270 Frogs of the Okefinokee Swamp

todes which were found in the alimentary tracts were Cosmocerella
haberi Steiner. Three other new species of nematodes were found in the
bodies of these frogs. They are to treated at greater length in an ap-
propriate journal elsewhere at some future date. (See Steiner, G., Some
Nemas from the Alimentary Tract of the Carolina Tree Frog (Hyla
carolinensis Pennant) Journ. Parasitology, Sept. 24, 1924, Vol. XI,
pp. 1-32).
7 per cent of these frogs were Opalina infested.

"In their feeding habits frogs of this species have tendencies which are of
beneficial economic importance. Rather frequently complaints are registered
concerning the presence of certain insect enemies of florists in greenhouses,
etc. Doubtless several dozens of the species of smaller tree frogs which
are common to a locality, if kept in such places would materially aid in keep-
ing down plant-lice, certain kinds of moths, beetles, mites, ants, cockroaches,
crickets, sow-bugs, diplopods, centipedes, etc. By means of adhesive discs
upon the tips of their phalanges, frogs of this species are able to adhere to the
smoothest kinds of vegetation and even to glass. They would damage few
if any of the most delicate of plants as they leap to and fro among the foliage
and ordinarily they absolutely would not damage any part of a plant by di-
rectly feeding upon its tissues."

ENEMIES

In 19 1 2 this tree frog was found in the stomach contents of only one
species of snake, the black snake. One had taken four of them, head first
and the long slender legs were the only undigested remains. One of the four
is a ripe female and doubtless this individual with the other three were cap-
tured while bound for the breeding ponds or while in them. The wonder is
that this bright common morsel is not more frequently captured, for the pine
woods tree frog (H. femoralis) was taken by four different species of snakes.

In 192 1 one of the Lee boys told me they caught "trout" {Micropterus
salmoides) with Hyla cinerea as bait. Doubtless on the prairies and along the
water courses when the Hyla cinerea descend to the waterlily level largemouth
bass and other fish feed on them as do water snakes and ribbon snakes.

On May 11, 1921, along the edge of Billy's Lake we thought that we ob-
served gnats, mosquitoes, etc., and other Diptera all over these frogs. All
the specimens we collected (however) according to the Cornell Department of
Entomology are of one kind, a harmless species (Oscinis longipes). These
frogs are convenient resting places, like the immediate vegetation in which
these flies were equally abundant. No doubt harmful Diptera prey on the frogs.

One of the reputable lumber foreman told us about the following episode
which with variations has been told of more than one species of Bufo. While
smoking he was seated on a log in the cypress bay. After a time he flipped
his cigarette and the lighted stub landed near a green tree frog. It seized
the stub in its mouth and went through interesting antics to eject it.

AUTUMNAL DISAPPEARANCE
From none of the Cornell Expeditions of December and January have we
received this species. Our latest residence in the swamp has been in August

Plate XXVII
Pine wood's tree-frog ( Hyla femoraUs)

1. Male croaking, Billy Id., Ga. May i6, 1921. Flashlight.

2. Male resting on bit of bark, Billy Id., Ga. May 16, 1921.

3. Male croaking in cornfield, Billy Id., Ga. June 4, 1921. Flashlight.

4. Tadpole, Anna's Pond, Folkston, Ga. June 27, 1922. Lateral aspect.

5. Two males, Billy Id., Ga. May 18, 192 1. Lateral aspect, x 0.33.

6. Transformed frog, Billy Id., Ga. July 25, 1921. Lateral aspect, x 0.33.

7. Eggs, stained, Chesser Id., Ga. July 4, 1922. x 1.3.

8. Female and male, Chesser Id., Ga. July 3, 1922. Ventral aspect, x i.o.

9. Eggs, stained, showing loose jelly envelopes, Chesser Id., Ga. July 27, 1922.

Hyla cinerea 271

and September when the species was active. On October 9, 1906, Mr. A. G.
Hammar secured three one year olds at Jennings, La., and we have another
record of transformation in October, 1915.

AFFINITIES

Holbrook (1842, Vol. IV, pp. 121, 122) reviews this in detail. According
to him Linnaeus held it a form of Rana arborea and credited it to Europe as
well as America {Rana arborea var. B.) Laurenti called it a variety of the
European Hyla viridis. Gmelin considered it a variety of Rana arborea.
Schneider called it a variety of Calamita arborea. Daudin pronounced it a
distinct form as does Holbrook, 1842, and LeConte 1856. The latter writes
(1856, p. 428) "Our American animal has but little resemblance to the Euro-
pean, especially in form."

Like Hyla gratiosa, Hyla femoralis, Hyla versicolor, Hyla cinerea has
indefinite outer envelopes. In egg deposition it may be between Hyla
gratiosa and Hyla andersonii group and the surface surface film group of
Hyla versicolor and Hyla femoralis. In some ways we incline to a closer re-
lationship with the former group. In tadpole mouth parts we placed it with
the Hyla gratiosa, Hyla andersonii group with their short 3rd lower labial
tooth row in contrast to the long 3rd lower labial row of teeth in Hyla squir-
ella, Hyla femoralis, and Hyla versicolor. In tadpole coloration and general
appearance it is more suggestive of Hyla gratiosa although the crests are
quite different. Of the American Hylas it seems all in all nearer Hyla ander-
sonii and Hyla gratiosa and somewhat approaching Hyla squirella.

BIBLIOGRAPHY
1907 Brimley, C. S. Journ. Elisha Mitchell Soc, Dec. 1907, Vol. XXIII, No. 4, pp.
158, 159-

1910 . Proc. Biol. Soc. Wash., Mar. 23, 1910, Vol. XXIII, p. 11.

1803 Daudin. Histoire Naturelle, Generale et Particuliere des Reptiles, Vol. VIII, p. 29.

1914 Deckert, R. F. Copeia No. 3, Feb. 14, 1914, p. 3.

1915 . Copeia No. 18, May 15, 1915, p. 4.

1842 DeKay, J. E. Natural History of New York. Zoology of New York, Vol. Ill, Part

III, p. 72.
1906 Dickerson, M. C. Frog Book, N. Y., p. 127.

1918 Dunn, E. R. Copeia, Jan 25, 1918, No. 53, p. 21.

1890-92 Garman, H. Bull. 111. State Lab. Nat. Hist., Vol. Ill, pp. 189, 346-348.

1915 Gronberger, S. M. Copeia, Nov. 19, 1915, No. 24, p. 55.

1926 Haber, Vernon R. Journal of Comparative Psychology, Apr. 1926, Vol. VI. No. 2,

pp. 214-216.
1856 Hallowell, E. Proc. Acad, of Nat. Sci. Phila., 1856, pp. 307, 308.
1825 LeConte, J. Annals Lye. Nat. Hist. N. Y. Vol. I, Part Second, 1825, p. 279.

1856 . Proc. Acad. Nat. Sci. Phila., 1856, p. 428.

1899 Miller, G. S., Jr. Proc. Biol. Soc. Wash., Sept. 28, 1899, Vol. XIII, pp. 75-78.
1924 Myers, G. S. Copeia, No. 131, June 30, 1924, p. 60.

1919 Pope, Philip H. Copeia, No. 76, Dec. 31, 1919, pp. 95, 96.
1924 Schmidt, Karl P. Copeia, No. 132, July 15, 1924, p. 68.

1923 Stejneger, L. and T. Barbour. A Check List of North American Amphibians and

Reptiles. 2nd Edition, Cambridge, 1923, p. 30.
1923 Viosca, Percy, Jr. Copeia, Feb. i, 1923, No. 115, p. 39.
1923 Wright, A. H. The Anatomical Record. Jan. 20, 1923, Vol. 24, No. 6, p. 406.
1923 • Copeia No. 115, Feb. i, 1923, p. 34.

1923 • Ecology VII, No. I, January 1926, pp. 82, 83.

1924 Wright, A. H. and Wright, A. A. American Naturalist, Vol. LVIII, July-August
1924, p. 378.

272

Frogs of the Okefinokee Sivamp
Hyla femoralis Latreille

(PI. II, Fig. 8; V, Fig. 2; VII, Fig. 3; X, Fig. 4; XII, Fig. 13; XV, Fig. 3; XVII; XXVII;
XXVIII; Text Fig. I, 17)

COMMON NAMES

Pine Woods Tree Frog. Pine Woods Tree Toad. Pine Tree Toad. Pine
Tree Frog. "Scraper Frog." Femoral Hyla.

RANGE

Check list. "Type Locality: Carolina. Range: Carolinas to Florida and
west to Texas." Stejneger and Barbour Check List 1923, p. 30.

Supplementary records. In 1926 (Wright, 1926, p. 82) we considered Hyla
femoralis as one of the eleven Okefinokee frogs "which occur mainly in the
Sabalian region or Gulf strip of the Lower Austral region." Its restricted
range in North Carolina (Brimley, 1907, pp. 159) in Craven and New Hanover
Counties and near Wilmington (Myers, 1924, p. 60) N. C, in South Carolina
at Mt. Pleasant (Schmidt, 1924, p. 68) and Goose Creek and Cope's record
for Georgia (Cope, 1889, p. 372) from Riceborough around to Nashville and
Allapaha directly west to Waycross, Ga., also confirms this interpretation.

In 1922 on June 8 we did not hear it near Cheraw, S. C, when Hyla
cinerea, H. andersonii, Rana virgatipes, Bufo fovderi, Hyla versicolor were
calling, nor at Columbia, S. C, June 9, nor at Millen Sta., June 10 {Hyla
squirella, Hyla cinerea, Hyla gratiosa, Hyla versicolor calling). On June 11
from the Altamaha near Jesup through Screven to Waycross and from Way-
cross to Folkston we heard them in the piney flat woods. In February, 191 1,
J. C. Bradley took the species at Waynesville, Ga.

In 19 1 7 our route led through Raleigh, Spartansburg, Atlanta, Montgom-
ery to Mobile, Alabama. Not until we reached the Tombigbee river below
Jackson in Washington County did we record our first specimens of this
species. Then we came into Sabalian influences. In 1923 Viosca (1923, p.
36) records it from the Longleaf Pine Hills of Louisiana.

Little is known about this species and writers have assumed it not so
abundant as others. Brimley (1910, p. 12) speaks of it as "apparently less
common than cinerea and squirella." In the piney woods we have more notes
on it than any other Hylid (except Acris) it was so abundant. Like Hyla
gratiosa at times it betakes itself into the high pines and is hard to get, and
like Hyla gratiosa is a decidedly Sabalian or Gulf strip species.

Local Okefinokee records. In 191 2 we took it in the following Okefinokee
localities: Billy Island, Mixon's Hammock, Honey Island, Prairie-heads.
In 192 1 on Billy, Chesser, Floyd, Craven, Black- Jack Islands, the Pocket,
Craven Hammock, Billy's Bay, Honey Island Prairie, Suwannee Canal, St.
Mary's and Suwannee Rivers. In 1922 we secured it at Trader's Hill, Camp
Pinckney, Folkston, Starhng Branch, St. Mary's, Chesser Island and other
places in Georgia and at Callahan, Florida. It is essentially a pine barrens
species.

Hyla femoralis 273

GENERAL APPEARANCE

Cope (1889, p. 371) writes: "Body short, rather broad, and the entire ap-
pearance as to pattern of color and shape not very dissimilar from Hyla versi-
color, from which, however, it is readily distinguishable by the femoral yellow
spots; the dark postocular vitta, the absence of light spots under the eyes."

Brimley (1907, p. 158) characterizes it as follows: "Back . . . markings
do not form an X-shaped mark. Back of thigh with yellow spots or variega-
tions. No hght spot below eye. No yellow spots on sides."

Deckert (1915, p. 3) who has studied southeastern frogs more intimately
than most observers says this species "resembles our own gray tree frog, with
its rough skin and star-shaped dark patch on the back, but is smaller and more
slender."

We would consider this species a small species. Cope considers 35 mm,
above the average size and his largest is 39 mm. LeConte a century ago
gave I 1/2-1 3/4 inches as the adult range and it has been repeated for this
little understood form. It may reach 45 mm., as he says, but of the 140
specimens from 20 mm. upwards we have none over 40 mm., the average,
30 mm.

COLORATION OF SPIRIT SPECIMENS (1 9 1 2)

In spirits the upper parts are usually some shades of dark brown, rarely
light brown; in life the color of all our specimens was invariably gray or
rarely grayish brown. The under parts are white. The blotch between the
eyes is constant but seldom triangular. From either cephalic corner it sends
a bar to the anterior end of the upper eyelid. From the two caudal corners
of this interorbital spot two branches extend obliquely backwards to another
median spot just back of the shoulder region on the anterior back. In two
specimens these branches are connected with the anterior dorsal spot but not
with the interorbital spot (just as in Holbrook's figure. Vol. 4, fig. 3) as most
texts usually represent it. Obliquely backward from the anterior dorsal spot
runs a dark bar to the middle of each side. Frequently between these last
two bars there are two more which extend backwards from the anterior
dorsal spot along the posterior back. Sometimes one or both of these bars is
disconnected with the anterior dorsal spot or almost absent. Several isolated
irregular spots sometimes take their place or represent the more regular bars.

From the tip of the snout a dark line extends to the eye. From the eye to
the ear it is a band with two dark borders. Back of the ear the lower border
ceases and the upper one passes over the shoulder to the middle of the side.
Whenever this lower border extends farther back it is more or less broken.
In all cases it more or less demarcates the separation of the brown or gray of
the back from the white of the belly. Beneath this hne there may be some of
the dorsal color but very little. When this line breaks up back of the middle
of the side, it often produces a motthng, reticulation or vermiculation along
the groin. In some specimens the line may be faint or almost absent on the
snout or back of the shoulder. In most of the larger specimens there is a dark
line along the posterior edge of the forearm and anterior edge of the foreleg.

2 74 Frogs of the Okefinokee Swainp

Most of the specimens have the transverse bars of the fore and hind Umbs
rather indistinct. Some have the whole throat to the white breast fold dotted,
others have just the chin dotted or only its sides. In the males the whole
throat area is much darker and the chin much the lighter. The posterior edge
of the thigh has the dorsal body color with three to ten distinct circular or
elliptical yellowish or yellowish white spots unlike the thigh reticulations in
Hijla versicolor. Sometimes in the larger forms the spots are fewer and ir-
regular, cordate or comma-shaped due to the fusion of spots.

STRUCTURAL CHARACTERS (WRITTEN IN 1912)

Upper surface more or less smooth, occasionally with scattered granula-
tions; belly, under surface of thigh and breast strongly areolate, the throat
granulate (smooth in one), the pectoral or breast fold smooth or slightly
granulate, rarely strongly granulate. Third finger longest; fourth finger
sUghtly longer than the second; fourth toe longest and with two phalanges
free; the web does extend as a margin to the disc of the longest toe hut is not so
wide as in Hyla versicolor; tympanic fold present, rarely almost absent;
vomerine teeth between the posterior nares; tongue usually slightly notched
as Cope figures ('89, Fig. 94, 4) but 5 specimens have no notch, two being
rounded and three truncate; two of the notched specimens had the right lobe
much smaller; nostrils nearer tip of the snout than the eye.

MEASUREMENTS
(Recent Material)

Head to angle of mouth 1.14 (20 mm.)-i.25 (28 mm.cf)-i.i (28 mm. 9 )-
1.2 (36 mm.cf)-i.3i (36 mm.9)-i.47 (40 mm. 9) in width of head; head to
rear of tympanum i. 06-1. o-i.o-i. 09-1. 09-1. 11 in width of head; head to
angle of mouth 2.85-3.5-3.1-3.6-3.8-3.7 in length of body; head to rear of
tympanum 2.6-2.8-2.8-3.27-3.13-3.0 in length of body; snout .625-.88-.80-
.70-.66-.91 in first finger; snout . 875-1. lo-i.i-i.i-i. 0-1.08 in fourth finger;
snout .625-.66-.70-.80-.66-.83 in first toe; eye i. 33-1. 5-1. 66-1. 25-1. 5-1. 33
in snout; eye .66-.66-.83-.71-.62-.55 in tympanum; eye .83-1.33-1.33-.875-
1.3 3- 1. 2 1 in first finger; tympanum 3.5-4.5-3.4-4,0-4.4-4.8 in intertympanic
width; tympanum 2.0-2.25-2.0-1.6-2.4-2.4 in snout; internasal width i.o-
1.0-1.0-.83-1.16-1.2 in upper eyelid width; interorbital width .8-.66-.71-
.57-.75-.625 in internasal width; interorbital width 2.8-3.0-2.4-2.8-2.75-3.0
in intertympanic width.

Forelimb: forelimb 1.66-2. 0-1.86-2. 1-2. i-i. 81 in length of body; forelimb
2. 5-3. 2-2. 73-3. 1-3. 18-2. 7 in hind limb; first finger i. 6-1. 62-1. 5-2.0-1. 87-1. 36
in third finger; second finger i. 33-1. 44-1. 2-1. 4-1. 5-1. 25 in third finger;
second finger .63-.88-.80-.70-.80-.91 in first finger; third finger .75-.77-.66-
.85-.80-.93 in second toe; fourth finger 1.14-1. 3-1. i-i. 27-1. 25-1. 15 in third
finger; fourth finger .71-.60-.63-.72-.66-.77 in first toe; internasal width
1. 25-2.0-1. 6-1. 16-1. 33-2. 2 in first finger, internasal width 1.5-2.25-2.0-
1. 66-1. 66-2. 4 in second finger; internasal width 2.0-3.25-2.4-2.33-2.5-3.0
in third finger; internasal width 1.75-2. 5-2. 2-1. 83-2.0-2. 6 in fourth finger.

Hijla fcmoralis 275

Hindlimb. length i. 5-1. 5-1. 46-1. 5-1. 5-1. 5 in hind Hmb; tibia 1.8-1.9-
1.8-2. 1-2. 05-1. 9 in length; tibia 2.7-2.9-2.64-3.1-3.08-2.8 in hind limb; tibia
1.1-.90-.96-1.0-.97-1.05 in forelimb; tibia .68-.70-.64-.70-.80-.76 in hind
foot; first toe 1.2-1.6-1.14-1.5-1.5-1.4 in second toe; first toe 2.4-2,66-1.7-
2.25-2.37-2.0 in third toe; first toe 2.8-3.33-2.85-2.9-3.0-3.0 in fourth toe;
first toe 2. 2-2. 66-1. 71-2. 12-2. 37-1. 9 in fifth toe; second toe 2.0-1.6-4.5-
1. 5-1. 58-1. 4 in third toe; second toe 2.33-2.0-2.5-1.9-1.5-1.58-1.4 in third
toe; second toe 2.33-2.0-2.5-1.9-2.0-2. 14 in fourth toe; second toe 1.8-1.6-
1. 5-1. 4-1. 58-1. 35 in fifth toe; third toe 1.16-1. 25-1. 66-1. 27-1. 26-1. 5 in
fourth toe; third toe .93-1.0-1.0-.94-1.0-.95 in fifth toe; fourth toe i. 07-1. 15-
1.0-1.05-1.16-1.07 in hind foot; fourth toe i. 57-1. 45-1. 55-1. 5-1. 46-1. 4 in
tibia; fourth toe 1.7-1.3-1.5-1.5-1.41-1.47 in fore limb; fifth toe i. 2-1. 25-
1. 66-1. 34-1. 26-1. 6 in fourth toe;internasal width i. 2-1. 5-1. 4-1. 33-1. 33-2.0 in
first toe; internasal width 1.5-2. 5-1. 6-2. 0-2. 0-2. 8 in second toe; internasal
width 3.0-4.0-2.4-3.0-3.16-4.0 in third toe; internasal width 3.5-5.0-4.0-
3.8-4.0-6.0 in fourth toe; internasal width 2.75-4.0-2.2-2.83-3.16-3.8 in
fifth toe.

HABITAT

Holbrook (1842, Vol. IV, p. 128) writes: "This little animal lives in the
deep forests of Carolina and Georgia; it chooses trees for its residence, and is
sometimes found thirty feet from the ground, feeding on such insects as choose
the same localities." In 1855 LeConte (p. 429) notes that they are not about
dwellings like Hyla squireUa. Deckert (1914, p. 3) in his "Listof Salientia f rom
near Jacksonville, Florida," (1911-1912) says " Hyla femoralis Latreille, (is)
common on pine trees, but not easily caught." In 19 15 (p. 3) the same author
adds: " Hijla femoralis Latreille is called the Pine tree toad, from its habit of
frequenting the tops of pine trees almost exclusively during the summer
months."

In 1925 Viosca in discussing the Florida Parishes or uplands of Louisiana
comments that "To the east of the Shortleaf Pine area are the Longleaf Pine
Hills with gentle slopes, and intertwined by winding creeks. Here we have a
limited representation of practically all species found in the uplands generally
and in addition, some species which may be said to be characteristic. Hyla
femoralis is the typical tree frog of this section."

At Leroy, Ala., June 12, 19 17, Dr. H. H. Knight while sweeping bushes
and lower branches of trees with an insect net for Caspids caught two young
Hyla femoralis.

In 19 1 2 we make the following notes on habitat: "On the outskirts of the
swamp we found this form in the cut-over lands. Here they were taken on
the small shrubs and on the ground. On the lily-covered prairies south of
Honey Island they were not infrequently found as was Hyla cinerea. A few
were observed on the Hly pads or in bushes of the little islets of this area.
Many of the specimens from Billy's Island were taken from the various build-
ings of the Lee family and particularly from the sides of the old cabin camp
of our party or in the small shrubs or trees nearby. Holbrook's observations
that it (Vol. 4, p. 128) differs from H. squirella "for it is never found near

276 Frogs of the Okefinokee Swamp

outhouses or about fences and in old fields" must consequently not be taken
too literally, though the conditions on Billy's Island even in Lee's clearing
are yet essentially forest-like in their nature.

In 192 1 we have made the following notes on habitat. On April 27 we
have "This morning four Hyla femoralis are dead. Two inches of rain water
in their can. Apparently they like water in general less than some other
forms and are rather delicate." On May 9 "Harry Chesser brought us a Hyla
femoralis which he caught on a cypress tree in a cypress bay. Surely it is not
restricted to pines." On May 7 we have a Hyla femoralis from the hammock,

FIRST APPEARANCE

In 1 92 1 we entered the swamp April 21. At once the boys began to bring

us pinewood's tree frogs. One specimen of April 25 was a spent female which

shows the specimen must have been abroad some time before. The earhest

seasonal record we have is a specimen taken February 19,1911, by Professor

. J. C. Bradley at Waynesville, Ga.

GENERAL HABITS

Metachrosis. In 1826 (p. 342) Harlan holds "its colors various: a variety
with the back chiefly occupied with a large irregular blotch; legs barred." In
1855 LeConte (p. 428) notes that "The darker marks in the head and back
are sometimes evanescent but the yellow spots on the hind part of the thighs
are always more or less distinctly visible. The chin is cancellately reticulate,
sometimes speckled with dusky, sometimes altogether of that color. The
color of the body above varies at the will of the animal."

Deckert (191 5, p. 3) writes that "Its usual color is dark reddish-brown or
chestnut, but it can readily change from fleshy pink or pale gray to any shade
of brown, gray or green."

In 192 1 a little 20 mm. Hyla femoralis has these colors (Non-Ridgway)
April 26: "Back greenish gray with dusky. Black line from snout through
eye over tympanum past angle of mouth half way to unguinal region. Upper
margin silvery or yellowish silvery; lower margin of black vitta yellow. There
is some pinkish bronze on rear end of vitta. Bright silvery or yellowish stripe
on back of hind foot, fainter on back of tibia. Yellow over vent and on to
femur posterior half way to knee. Only three rear orange yellow spots on
each femur."

On April 27, 192 1 we have the note "James Lee says that he found on rail
fence some which are whiter than those he finds in the rain barrels which have
boards or cover over the top." One rain barrel specimen we watched for
several days before May 2. On May 4 we observed that it "had remained
several days. It is as dark as the wood over it." On the following day. May
5, it was still in the barrel.

Two weeks later. May 19, "on the way back from a trip to Crosby Pond
at 6:00 p. m. near the remains of an old cypress pond in piney woods saw a
female Hyla femoralis hopping along into saw palmetto. It was as whitish

Hyla Jemoralis 277

gray as any Hyla versicolor I ever saw. The spot in middle of back showed
beautifully, also spot between the eyes. This female I took out to look at it.
It leaped away on to gray sand. Has a hard time seeing it, it matched the
gray sand so well. In one minute since its capture it had darkened consider-
ably. In denser cover 9 inches high with small saw palmetto, Vaccinium
myrsinites, calico bush, Gaylussacia frondosa, etc., found a half grown H.
femoralis. It was green on its back (very suggestive in size, color and appear-
ance or marks of Hyla squirella, which strangely enough we don't get here).
A little later lost another Hyla femoralis in same place as the green one.
Anent the half grown green specimen on May 21 we have the note: "In the
compartment of Hyla jemoralis, most of the specimens, in fact all (including
one little half grown one green when captured) are Vandyke brown or moss
brown." "A captive female in jar June 19 is pale hght mouse gray on back
with no markings revealed." On July 18, Harper makes the note: ^' Hyla
femoralis on green palmetto frond in sun in p.b. about 4:00 p. m., bleached
out, much as at night; bending the neck back, to rival Chorophilus.'^

General habits. In 192 1 we acquired the following notes on their daytime,
non-breeding habits and resting.

On April 23, the boys found two on the rail fence at 2 :3o p. m. The next
day they brought three more from the same fence. On April 26, the boys
found some more Hyla femoralis in the rain barrels along the railroad and
near the company's wood pile. "In a pine near camp about 15 or 20 feet up
on the end fork of large branch is a Hyla femoralis male. It doubtless is the
one we have heard ever since we have been here." On April 27 the boys
brought in another male from the rain barrels along the railroad. The rain
barrels too are used in putting out the trees when they start to burn.

VOICE

LeConte, 1855, seems to be the first to note that it differed from H.
squirella in voice. No one since has remarked about its voice until Deckert
(1915, pp. 3, 4) writes that "The noise resulting from the calls of the males on
these occasions, is deafening. This call cannot be reproduced on paper, being
a rapid succession of harsh, rattling notes, higher in pitch than the call of H.
squirella, and kept up all night. During the dry season this tree toad occasion-
ally calls from the tops of the pine trees, one answering the other."

In 192 1 the first general calling of Hyla femoralis came May 14. Then the
author "Went to Billy's Lake Landing. Worked eastward. Heard plenty of
Hyla femoralis in trees. They are approaching or are on the edge of cypress
ponds. Some are yet high in the trees. They are calling at intervals all over
the piney woods, particularly near the edges of cypress bays. One call, an
interval, one very faint call, interval. ..."

On May 16 "When a threatening storm passed over Hyla femoralis from
the trees were almost in chorus. Around cypress ponds several calling. One
was on a projecting piece of pine bark on the tree and within reach, about six
feet from the ground. Harper saw it croak. Afterwards the throat pulsated
all the time." Harper notes that it "let me come up in plain sight, within a

278 Frogs of the Okefinokee Swamp

couple of yards and croaked for me on its bark slab-perch. Throat kept dis-
tended while its sides, or rather whole body, vibrate. Kek, kek about 20 times,
usually ending in krak, krak. Surely a cracked voice." Later in the evening
we visited this croaker but "it was gone and not on the bit of projecting bark.
Soon we understood. In two or three trees low down we heard Hylafemoralis.
Their notes are speeded up, more extended and continuous than when high
in the pines. One we found on the moss. The instant we' entered (another)
pond we heard a queer note and it was from the trunk of a bay tree right
near us. It proved to be a male Hyla femoralis. All around us they were
calling. One was in a bush 3 feet above ground. Another on moist ground at
edge of pond in amongst six foot sedges. When a male croaks it is the lower
throat which swells out, not the chin region as well. Brought back three
Hyla femoralis."

On May 21 "After the rain we started for the turpentine still. In piney
woods heard and saw Hyla femoralis all afternoon but isolated calls. When a
warm shower came they leaped into chorus. Sounds like a peculiar Cicada
note in chorus, only once in a while is the frog-like character revealed when a
frog ends its call with the last two or three squeaks. This gives it away. The
chorus is one continuous stridulating din and goes down the piney woods like
a wave. Rain is not over. They are at it yet (5:30 p. m.)."

During a congress when several species are breeding in the same pond the
machine gun calls of Hylafemoralis make it difficult to hear or time other calls.
Once when we were timing the intervals in Gastrophyne' s calls Hyla femoralis
calls would break in frequently. In 192 1 and 1922 we recorded several in-
stances where their calls drowned out the calls of Hyla gratiosa, Hyla squirella,
Bufo quercicus and Pseudacris ocularis. We tried the experiment of half
closing our ears to close out the Hyla femoralis calls. The other sounds came
out very distinctly in the attempt. The breeding evening congress call is the
diurnal Kek speeded up, as many as 6 or 7 coming in a second. We have
counted 60 or 70 or more calls in rapid succession without deflation.

1921

April 24. Heard Hylafemoralis in trees 7:30 p. m. Long Pond.

April 25. Near Long Pond one or two calling.

April 27. In the trees H. femoralis are calling in isolated cases but more than
heretofore. Day overcast. 67°. At 9:30 heard a few in trees.
Air 76°.

April 28. About three heard.

April 30. Tonight air cold. Temperatures range from 50-70°. Little hu-
midity, brisk. Over in Long Pond and elsewhere we hear no Hyla
femoralis. As yet we have not heard them at night.

May 3. Tonight at 8:00 air 56°. The last few days have been so cool that
we have heard no Hylafemoralis.

May 9. Occasionally during day (air up to 84° or higher) hear Hylafemoralis.

May II. Temperature 7o°F. at 6:30 a. m. Feels Hke rain. Hear several
Hyla femoralis.

May 12. Considerable rain. One Hyla femoralis heard.

May 14. At 7:00 p. m. several Hyla femoralis calling. During day several
calling from pine trees.

Hyla femoralis

279

Mi

May

15-

16.

May

17-

May

18.

Mav

19.

May

20.

May

21.

May

24.

May

27.

May

28.

May

30.

June

I.

June

2.

June

3-

June

4-

June

5-

June

12.

June

13-

June

14.

June

16.

June

18.

June

20.

June

23-

June

24.

June

25-

June

26.

June

28.

June

29.

July

I.

July

3-

July

6.

July

7.

July

9-

July

10.

July

II.

July

14.

July

15-

July

16.

July

17-

July

23'

July

24.

July

27.

July

28.

July

29.

August

9-

August

10.

August

14.

August

15-

August

16.

August

22,

Rains from east all day. Hyla femoralis calling at night. Tem-
perature, today 66-69°; tonight 66-67°. During day once in a while
Hyla femoralis calls.

Storm threatening, brought almost a chorus in pine trees today.
Very active at night.

Heard Hyla femoralis several times but not very active.
Chorus at 9:30 p. m.
Several calling around edges of ponds.

One calling. We have heard this species soon after camp was es-
tablished. Several calling in woods.
No end of Hyla femoralis calling at night. Air 70°.
Tonight very few calling 10:00 p. m. Air 77°. A dry non-humid
day.

Hear Hyla femoralis about camp. Air 78°, 1:20 p. m.
Hear one at 8:00 a. m. Air 82°.
Hear several 7 :oo a. m.

On Jones Island hear Hyla femoralis. Common near canal.
Two heard.

None calling on Island. Common on Billy Island Bay.
Hear 7:20 a. m., pinewood tree frog in trees. Sky overcast. Air 73°.
No end of Hyla femoralis calling and abroad.
One heard in hammock.
One or two heard.
One heard. Air 71°.
Heard 7:00 a. m., one.
One heard 6:15 p. m.

From time to time heard quite a few. Air 75°.
After two inches rain no end of them in chorus.
Males calling.
Plenty calling.
Heard a few.
Calling commonly.
Common.
No end at night.
Great chorus. Air about 70°.
Heard one.

Heard one or two after dark.
One calling at 8:45 p. m.
Rain. Great chorus.
None calling.

One or two 6:30 a. m. Air 74°.
Heard them.

Abundant at Coatbet Pond 8:30-10:00 p. m.
Calling lustily.
Some calling at 7:45 a. m.

Hyla femoralis common at night. Several at 8 :oo a. m. and 1 2 noon.
As night comes on several start up. Black Jack Island.
Abundantly calling.
Common.

Calling along St. Mary's River.
Several calling.
Several.
Two heard.
Commonly heard.
Hijla femoralis at Billy's Island camp.

Frogs of the Okefinokee Swamp

June

12.

June

13-

June

14.

June

18.

June

22.

June

23-

June

25-

June

26.

June

28.

June

29.

July

2.

July

3-

July

4-

July

8.

July

9-

July

10.

July

II.

July

12.

July

13-

July

IS-

July

16.

July

17-

July

18.

July

19.

July

21.

July

23-

July

25-

July

26.

July

28.

July

29.

July

30-

August

I.

August

2.

August

4-

August

5-

August

8.

August

9-

August

10.

August

II.

August

12.

August

13-

August

14.

August

15-

August

16.

August

ly.

August

18.

August

19.

August

20,

August

26.

August

30.

1922

Chesser Island, few in corn field calling at dark.

Few calling in hammock.

Several around Coat Bet Pond at night. Few heard around camp

during day.

Heard one in hammock at i :oo p. m.

Heard several around camp 6:30 p. m.

Heard a few pines around open pond near Trader's Hill.

Two in hammock at camp.

Several calling after sunset.

Slight rain. Several heard in hammock.

Heard a few in oaks 8:05 a. m.

Heard many at Starling Branch 8:30 p. m. -midnight.

Rainy 1-3:00 p. m. Hyla femoralis calling some.

One heard.

Several heard in hammock during shower.

One heard at Thompson Landing, St. Mary's River.

Common in cypress bay.

Several heard during the day.

Around edge of Coat Bet many calling. A large assembly.

One heard at 10:00 a. m. in pine barrens.

A few heard.

Heard at 7:40 a. m.

Several calling 6-7 p. m.

Several heard.

Common in rain.

Many late in evening.

A few at Coatbet Pond.

Quite a few calling but not as many as we would expect after a

hard rain.

No end of calling males. Air 86°.

One heard on prairie 9:00 a. m.

One heard at camp.

Several heard at 7 :oo a. m.

Waycross, Ga. Hyla femoralis numerous along Dixie Highway at

night.

Hopkins, Ga. One heard 11:15 ^- "^•

Several calling in rain p. m.

Several calling 7:30 a. m. About 7o°F.

Big chorus at night. Temperature 79°. After today's rain.

Commonly calling after dark.

H. femoralis quite numerous.

Heard a very few at Camp Pinckney — Chesser Island 8:00 p. m.

to 12:00 midnight.

Several at nigh-t.

Commonly heard often in chorus 7-10:30 p. m.

A very few heard.

One heard.

Common in Spadefoot ponds.

Strong chorus in evening.

One heard during evening.

Congress at night.

Common along road near St. George.

One at 9:00 a. m. in hammock.

Several heard.

Hyla fern or a lis 281

Sept. 9. Heard at 6:20 p. m.
Sept. 12. Heard at 6:40 p. m.
Sept. 16. Several heard at night during heavy rain.

In 192 1 in the last of April and first of May we had a good index of Hyla
femoralis activity. On April 24, April 25 and April 27 under influence of
65-75° temperatures and cloudy weather we heard a few, but on April 30 to
May 8 minima range from 40-54°, average 48 or below, maxima from 73-83,
average 77 for four stations around the swamp. The cold, brisk, clear weather
kept them inactive and none were calling. There was no or little rain before
May 1 1 when several Hyla femoralis began to call. The air felt like rain on
this date and for two weeks considerable rain fell. Hyla femoralis became
very active. Minima 60-70, maxima 73-89, average 82. It, therefore, was
not the maxima of April 30-May 8 or May 1 1-25 which played the prominent
part but the higher minima 60-70 (not 40-54 of April 30-May 8) and most
of all humidity were the important factors. These periods are typical and
no need of reviewing the whole period of 192 1 and 1922 in necessary.

MATING

Male (From life, June 5, 1921). Cephalic half of pectoral fold pale
vinaceous drab to darkish grayish brown, posterior half like the belly. Throat
darker than the belly.

Female (From life, June 5, 1921). General color sorghum brown or deep
brownish drab or mars brown on back. Black spot between eyes. Another
spot with four points two behind and two ahead, the cephaHc ends above
tympanum. Two on either side of the middle of the back and one over the
crupper. Black or deep brown line from snout through nostril to eye narrow.
From eye through tympanum to groin where it breaks up into spots. Another
of same color on back of fore foot, fore arm, front of fore leg and over vent.
Below white line of the vent is a black one. Spots on rear of thighs orange to
chrome or light cadmium. One female with grayish white spots instead of
orange spots. Under parts white. Pectoral fold pure white to angle of the
mouth. Chin white with fine black spots.

Iris ecru drab, drab gray, pale vinaceous drab with reticulation of black in it.

Structural differences. In the Hylidae as Bufonidae we are accustomed to
saying that the throat of the female is not dark but more or less like the rest
of the venter. In many of the females of Hyla femoralis the throat is white
but sometimes the prominent white granulations have brown reticulations
around their bases. Occasionally only the tip of the granulations is white and
the general effect of the throat is very dark.

The measurements of 134 adults, 70 males and 64 females yield the
following results: males range 24-37 mm., mode 31 mm., average 30 mm.;
females, range 23-40 mm., mode 36 mm., average 32 mm. A study of the
modes for these sexes reveals the following: The males and females start about
24 mm. The first mode for the males comes at 28 (9 specimens) mm. while
the first female mode is 30 (9 specimens). The second mode for the males is

282 Frogs of the Okefinokee Swamp

31 mm. (17 specimens) while the second for females is 33 (8 specimens). The
males then taper from 3 1 mm., down to 37 mm. while the females have another
mode at 36 of 10 specimens. The males thus seem to be 2 mm. more or less
smaller at each mode and at the largest size than the female. The largest
male is 37 while we have two females 40 mm. The bulk of the breedirg males
and females came from 27 mm. onwards or at the beginning of the second year.
The few breeding males and females at 24, 25 and 26 mm. are doubtless also
at the beginning of the second year rather than at the end of the first year.

Duration, night or day. On the night of May 18, 192 1, at 9:30 p. m. at
Long Pond "There was a perfect din in cypress bays and at the north edge
of this pond from Hyla femoralis. Some were in bushes, other on bay trunks,
and others were hopping in moss edges or grassy edges of the pond or amongst
the lizard's-tail (Saururus). Could find no mated pairs. On the moss edge,
not in pond proper found Hyla cinerea pair." On the following day. May 19
"the boys at different times brought in females of Hyla femoralis found on
the ground in different parts of the island. They must be bound for the ponds.
We found one in the hammock on the ground near Long Pond (see above
May 18)."

On May 21, 192 1 after 8:00 p. m. we found a congress of Hyla femoralis
and these are our rough notes: "We found a Hyla femoralis on a log croaking;
one or several on the grass; many in trees and bushes 2-4 or 6 feet above
ground or water. One male is croaking and crawling up the tree right after
another male which is not croaking. The males after a time might dislike the
flashhght and walk up the tree for a foot more or less. We found some
croaking in amongst some pine bush in the water. Harrison Lee espied a
pair mated on a branch of the pine brush. The pair i-i 1/2 feet above water.
We flashed them twice. They did not move. Broke when I captured them.
Put them together in a jar. Soon they were mated again. Brought them to
camp at 1 1 130 p. m. At 1 2 :oo midnight put in a porcelain pan for photography.
In the morning at 6:00 they had laid and broken." The following night little
happened. "One male was calling from bush beside Newt Pond. On the
pond no end of Hyla femoroUs eggs. The males crawl up under grass edges or
logs or any cover. Are they more or less subterranean?"

Another congress of June 4, 192 1, reveals other points of interest: "No
end of Hyla femoralis. Where do they come from? On one palmetto within
2 or 3 feet of each other were 5 male Hyla femoralis calling. Later Marion
Lee found a Hyla femoralis pair on saw palmetto leaf i 1/2 feet above ground.
Took another pair on the moss at the edge of pond. Everywhere in the road
home were Hyla femoralis migrating. In little transient pools no end of Hyla
femoralis. Took another pair Hyla femoralis moving along in the path.
Tonight frogs are laying in all sorts of transient places.

In 1922 we made these two pertinent series of journal observations. On
"July 3, 8:00 p. m.-i :oo a. m. July 4 in a pond 1/2 mile west of Trader's Hill,
Ga., there was a great din of these frogs in a shallow grassy pond. No end of
Bufo quercicus, a few Hyla squirella, 4 Hyla gratiosa calling, a few toads,
plenty of Acris. Never heard such a composite racket, and our ears rang for

Hyla femoralis 283

a long period afterwards. Took no end of Hyla femoralis males. Took a
pair in the grassy water. At Anna's Pond the Hyla femoralis were common
in the saw palmettoes around the edge of the open pond. Equally common
were Hyla squirella which frequent the same place. The call of Hyla femoralis
is much more rapid than that of Hyla squirella. Caught a pair of Hyla
femoralis at this pond."

On July II at night "7 :3o-ii :oo p. m. we hear no end of Hyla femoralis in
every crossmg and cypress pond. At the second 'dreen' we found many of
them. They looked very queer as they skipped along the water ahead of the
auto hghts. They were especially centered about a clump of saw palmettoes.
Here Miles Pirnie found a pair and I another. Later on on one blade were
two more pairs and one male. In this clump were possibly 20 or 30 males
calling. This breeding came after a hard rain."

The females doubtless do not enter the ponds until about ready to lay.
On May 19, 192 1, "the boys brought in several females on ground hopping
along. They must be making for the ponds. Found one on the ground on
hammock near Long Pond." In one pond June 5, 192 1, after a breeding
congress "we found two dead female Hyla femoralis.'^ Doubtless they suffer
as do females at a Bnfo congress. All through May, June and July of 192 1
the boys on Billy's Island used to find females hiding under grass during the
day or hopping around on the island. In 1922 on Chesser Island, June 29,
the Chesser girls "caught a female travelling along on the ground. It was
gravid yet not swollen hke some species." The next day "they found another
female on the ground in the chufa field where they were working. It was in
the shade." A week later, July 6, "Mr. Chesser hoed out another female in
the open field. There has been a chorus of them in the pond nearby. Probably
the female was resting here during the day." Five days later, July 11, 1922,
we noted "at 10:00 a. m. a female crossing our path in the hammock headed
for the pond where in the evening we recorded a great congress." Until a
month later, August 6, we saw females hopping along in the hammock from
time to time. One on last date was found moving toward a pond at 6 :oo p. m.
At 7 :oo p. m. the chorus at this pond began in real earnest.

Amplexation (Normal, abnormal, cross). On May 21, 192 1, we found
the first pair mated in axillary fashion at about 11-11:30 p. m. They laid
before the following morning. Two pairs captured the night of June 4-5 laid
the same night and had axillary embrace. We found July 3, 192 1, two pairs
in overflowed sweet potato fields each normal in amplexation. So also another
pair in Newt Pond. "The last pair we wished to observe but a toad butted
themand we lost them." On July 2, 1922 between "8:30 p. m.-i 2:00 midnight
caught a pair between i :oo and 4:00 a. m. Same night they laid their com-
plement. On the night of July 3, 8:00 p. m.-i:oo a. m., July 4, secured two
pairs each at a different pond. Like the predecessors they embraced axillary-
like and each pair laid eggs the same night between i :oo-7 :oo a. m. On July
II, 1922 between 7 :3o-i i :oo p. m. we secured four pairs. Each was normal in
amplexation and laid this same night. We recorded no cross embraces in this
species. We have this note of males mating for May 21, 192 1. "Some of the

284 Frogs of the Okefinokee Swamp

males caught last night are now mated. These are axillary as was the mated
pair." On June 4, 192 1, we noted that "One of the Hyla femoralis pairs has
the male larger than the female. Its left arm comes on to the breast to the
middle hne and the other hand is in the axilla or a little lower."

OVULATION

Miss M. C. Dickerson (1906, p. 151) writes of it that "Nothing is on record
regarding the life history or habits." In 19 15 Deckert (p. 3) calls it "one of
the shyest of tree toads, rarely caught except during the breeding season, when
it frequents certain 'bayous' in thousands."

Habitat. We found breeding congresses in grassy transient pools near the
roads in the piney woods, in open ponds in cutover roads, in pools or ditches
beside the railroad and roads, in cypress ponds and in cypress bays. We found
mated pairs in overflowed grassy fields, shallow transient depressions, in
temporary overflows or drenched cultivated fields, swamps or dreens in the
cypress bays. Tadpoles were taken in pools beside Indian mounds, railroads,
roads, in cypress ponds and sometimes on the prairies, in diverse ponds on the
east mainland or in bays outside the swamp.

Period. In 1920 we (1920, p. 29) held that "The . . . tree frogs, like the
. . . pine wood's tree frog, lay their eggs from the very last of May to July."
Three years later (1923, p. 34) we considered Hyla femoralis as one of the five
which start breeding from May 15 on to June i. The next year we (1924, p.
37) made the "Season May 16 to August 21, for Hyla femoralis."

In 19 1 2 we made the following notes: "The breeding season must be about
the same period as for Hyla carolinensis. The two specimens taken (May
6-23) by Mr. Harper were male and female respectively. The latter was full
of ripe eggs. One spent female was captured June 5 and another June 18.
From May 30- June 22, numerous gravid females were observed or taken. Far
more than half of the adults were females and all of the specimens (11 in all)
from 33-37.5 mm., in length were of this sex while the largest male proved
32 mm. In the males the skin of the throat region seems more loose and dark
in color as in Hyla versicolor males. This large number of females seems
indicative of the approach or arrival of the breeding period. The wonder is
that none of us ever remembers hearing this form during the day, yet we
quite frequently found the species active and the examination of the females
showed them at the breeding point. Usually at the height of such periods
Hylas call by day as well as by night. No doubt some of the frog chorus of
the evening was of this species though we did not identify its call." This
implies May 6- June 22.

On the basis of gravid or spent females we have in 19 12 records of spent
females, on May 29, June 5, June 7, June 15, June 16, June 18-20 (2), June
25-27. Of gravid females we recorded 13 frogs as follows: One before May 28,
May 30- June 2 (3), June i, June 5 (2), June 6-7 (2), June 12 (2), June 18-20,
June 21 (2). This means they begin breeding before May 28, and continued
after June 21. In 192 1 on the same basis we have spent females April 25 (4),
May 3, May 17, May 16-23, May 30 (2), June 5, June 23, June 30, July 3.

Hyla femoralis 285

Gravid females: April 25, April 26, June 3, June 4 (3), June 9 (2), July 3.
This gives a period beginning before April- July 3. In 1922 we have spent
females from July 3-August 7 and gravid females June 29, July 2, July 3 (3),
July 6, July 11, July 12. This gives a restricted period of June 29- July 12.

On the basis of mated pairs we have the first date May 20, 192 1, followed
by records on May 21, 1921, June 4, 1921, July 3, 192 1, July 2, 1922, July 3,
1922 (2), July II, 1922 (4), or a period from May 20- July 11. On the basis of
recorded field eggs we have eggs taken May 21, 192 1, May 22, 192 1, May 26,
1921, May 27, 1921, June 4, 1921, June 5, 1921, June 6, 1921, July 3, 1921.
In 1922 we have field eggs July 3, July 4, July 11, July 13, July 27, August 9,
August 17. In 1922, the last chorus was recorded August 20, though some
called until mid-September.

Viewed from records of spent or unspent females, choruses, field egg rec-
ords, mated pairs and transformation dates (June i6-August 9) the ovulation
begins as early as April 20 or 25 and extends to late August or September i.

Temperature and humidity. In 192 1 the temperature of ovulation periods
were 82-95 maxima and minima 61-74°. Either before, during or after these
periods rains of 1.22, 1.60, 2.80, 1.76, i.oo, 2.88, 2.27 and 1.60 inches prevailed
in stations around the swamp. In 1922 the maxima were 81-96°, the minima
64-74°, the rains before, during or after breeding were 2.00, .80, .48, 1.80, 1.60
and 3.47 inches in stations about the swamp. With this species, like other
southeastern species, humidity is the greatest factor. Temperature is sec-
ondary if it be above a certain essential minimum.

Egg-laying process. The films are so like those of Hyla versicolor they must
lay their eggs in the same fashion. We often found groups of films 3-8 in
number. These groups doubtless represented the 500-768 complement of one
female. In the first captive pair the one film had 5 centers. The large films
of 200, 150, 125 or 100 would not yield more than 3 or 4 films for a complement
(500 eggs) if each film of the complement were large.

EGGS

Attachment, egg mass. The first mass identified positively came the night
of May 20-21, 192 1, for a pair captured that evening. This was laid in cap-
tivity. "The mass is 15-16 inches long. All connected, this is partially
floating and partially resting on the bottom of the tray. The water is 3/4 of
an inch deep. This floating mass suggests the same of Hyla versicolor. There
are five centers of oviposition apparent." Our first field notes at the same
time are: "Found egg band attached to stems of grasses. It is 2-3 inches long
and 1/2 inch in width, brown above and yellowish or cream below. Now it is
1/4-1/2 or I inch below surface of the water. Was it at surface before today's
rain or laid beneath surface? This pond is out in the open. Another comple-
ment of three bands; one 3-4 inches long; another 6 inches; the third 4-5 inches,
brown and yellowish. This is in water below the surface slightly and amongst
grass where two small pine top branches had faflen in the water. Find another
complement of four masses one 2x2 inches; one 1x1/2 inches in diameter;
one bandlike. Probably close to surface before today's rain. Place where

286 Frogs of the Okefinokee Swamp

last two complements are found is directly under the pine brush on which we
caught pair last night."

On the following day, May 22 "found an area at the end of pond amongst
sticks, trash and chips where eight film packets of Hyla femoralis eggs were
laid. In the railroad ditches with oil and dirt on the surface the masses stand
out as black areas darker than the rest of the surface of the water. In another
part of the ditch three films of Hyla femoralis with a Gastrophryne egg film
4x7 inches. In another pond clear 20 x 3 feet found floating amongst
grassy edges a packet of 100-125 eggs. They are brown and yellowish. One
mass 6x3 inches separated from another round mass 2 1/2x2 1/2 inches.
More eggs in a mass than in Hyla versicolor. Insects get into fresh masses
and get stuck. Some masses amongst roots. Two masses either side of a
floating root, one of them 3 x :i, another 3x2 inches.

On May 27, 1921, in a run found numerous spoiled R. sphenocephala egg
masses beneath the surface. Above these eggs on the surface were Hyla
femoralis films. The surface temperature is 85-90°. One film floating free on
water. One film has 200 eggs, another 150 eggs, another 125 and the rest
smaller. Film may be eUiptical, circular and otherwise. Films floating some
distance. Have seen them go 10 feet in as many minutes. Some films at-
tached to ends of iris leaves. First eggs we recorded were in bands but films
are normal. If one touches the surface of a film the part touched adheres to
the finger. If loosened the part falls back in a little ball, reveals the yellowish
vegetative pole or it will cause the film to take a band shape first recorded for
this species. Probably the films were fastened or stuck to grass and rain
raised them and gave them the band-like shape as a consequence. One film
laid on moss in water's edge."

On June 5, 192 1, "in one pond south of our old 191 2 camp were no end of
Hyla femoralis eggs at the water's surface. The litter had all drifted to the
masses. Water surface between masses clear. Jelly just covered with trash.
Easy to tell where mass was and where not."

In 1922, we have a few additional pertinent notes. Of a mass laid by a
captive pair July 4 there is the note: "Eggs are in a loose string. Outer
envelopes merge into an outer mass Hke Ambystoma masses. Inner envelopes
with vitelH show distinct in the jelly mass. Nevertheless this is a film. Hyla
squirella does not seem to have this appearance. In another pond were many
films which are doubtless Hyla femoralis eggs. There are some quite band-
like. Are any of these Hyla squirella eggs?" On July 27, in a pond covered
with spoiled eggs was one fresh mass on the surface 2 x 14 inches in length.
The mass was wavy. After a very heavy rain August 17 spadefoots filled a
pond with its eggs, every grass blade being bent over with them. All over the
surface were little packets of fresh Hyla femoralis eggs.

On the night of June 4-5, 192 1, a female laid its complement without an
attendant male.

Egg description. In 191 2 our data were as follows: "The ovarian eggs are
brown on the animal poles and creamy or yellowish white on the vegetative
poles. In fact they look very much like those of H. versicolor and H. cinerea.

Plate XXVIII
Pine wood's tree-frog ( Hyla femoralis)
Pool with egg clusters, Billy Id., Ga. May 22, 192 1.
Seven egg clusters, showing black on greasy surface of trackside pool, Billy Id.

Ga. May 22, 192 1.
Eggs, Billy Id., Ga. May 22, 1921.

Male croaking, Billy Id., Ga. May 16, 192 1. Flashlight.
Male croaking, Billy Id., Ga. June 29, 192 1. Flashlight.
6. 7. 8. Tadpoles, Anna's pond, Folkston, Ga. June 27, 1922. x i.e.
9. Tadpole with 4 legs, Anna's pond, Folkston, Ga. June 27, 1922. x i.o.
[o. Tadpole with shrinking tail, Chesser Id., Ga. July 13, 1922. x 1.0.
Tadpole with stub of tail, Chesser Id., Ga. July 6, 1922. x i.o.
Transformed frog, Chesser Id., Ga. July 2, 1922. Donsal aspect, x 1.0.

Hyla fe moralis 287

A count of the eggs in the right ovary of the largest ripe female (37.5 mm.
long) gives 273 eggs or approximately 500 or 550 for the complete complement.
This equals or exceeds the number to be found in the average female Carolina
tree frog. The latter may exceed the pine woods tree frog by an inch in
length but the extreme slenderness in the loins, even in gravid females does
not admit of a large egg complement as in the short stocky Hylafemoralis the
ripe females of which are discernible at once. The ovarian eggs of these two
species are practically the same in diameter and we thus have a relatively
larger complement in Hyla femoralis. Actually the ovarian eggs of Hyla
cinerea are slightly larger but this does not vitiate the result. The measure-
ments of ovarian eggs of the two species gives .8-1.2 mm. for those of Hyla
femoralis and 1.1-1.5 mm. for those of Hyla cinerea. In proportions, appear-
ance and habits this species is a miniature Hyla versicolor and no doubt its
breeding habits are very similar."

In 192 1 our field notes on eggs are: ''Eggs laid in captivity May 21, 192 1
have vitellus .8-.9 mm., inner envelope 1.4-1.6 mm.; outer envelope loose, not
distinct from 4-6 mm., sometimes as much as 8 mm. Most of eggs without
circular outer envelope distinct because connected with the mass. Small
edition of Hyla versicolor. Egg cream or yellowish on vegetative pole,
brownish on animal pole."

On June 4, 192 1, in comparing Hyla femoralis and Hyla cinerea ^^ Hyla
femoralis eggs are smaller, vitellus .8-9 mm., inner envelope 1.4-1.6, outer
envelope indeterminate. Vegetative pole quite yellowish or cream, more
yellowish than in Hyla cinerea. ^^

Laboratory examinations of 58 eggs reveal 18 with vitellus .8 mm., 7 with
vitellus .9 mm., 31, i.o mm. and 2, 1.2 mm., average .95 mm., mode i.o mm.
The inner envelopes ranged from i .4-2.0 mm. The outer envelope was seldom
visible, occasionally distinct, 4, 5 or 7 mm. in diameter.

On June 29, 1922, the count of the left ovary of a gravid female showed
384 eggs to be laid, a possible complement of 768 eggs. On the following day
another female's left ovary was counted. It gave 382 or a possible 766 in all.
The complement reaches from 500-768 eggs.

Dangers. This species lays in any pine barrens pool. Many a shallow
grassy pool will have packets of their eggs. Many are caught by hatching;
certainly many of the tadpoles never mature. There must be a great loss in
this species. On July 3, 192 1 , they were laying in the flooded furrows of cane>
corn and sweet potato fields. These were all lost. On June 4, 192 1, we made
the notes "Tonight the frogs of several species {Hyla femoralis included) are
laying in all sorts of transient places."

In 1922 the species bred in great numbers in a crossway on the night of
July II. On July 13 the cross-way was practically dry. "All the Hyla
femoralis eggs found July 11 were now gone if they ever hatched." On July
27 in one pond we found "all over the pond spoiled eggs of Hyla femoralis
caught no doubt by a decided change of level."

288 Frogs of the Okefinokee Swamp

HATCHING PERIOD

"Eggs laid early morning of May 21, 192 1, hatched early morning of May
24, 1921, or three days later." Eggs "collected in the field in the morning of
May 22 after the laying of the night of May 20-21 hatched the night of May
23-24" or three days later. Eggs "laid the night of June 4-5, 192 1, after mid-
night were hatched by June 7, 6:00 a. m. or three days after ovulation."
The eggs laid by one captive pair July 4, 192 1, did not hatch at all. Doubtless
captivity, infertility, restricted quarters or laying without male may have
caused it.

MATURE TADPOLE

Color description from life (July 8, July 16, 1921). Upper parts of body
olivaceous black to dull greenish black. Throat on either side below eye
pomegranate purple. Belly on sides and across breast light vinaceous purple;
center of belly solid sulphur-yellow. Chin clear with very little or no black
dots. In younger tadpoles general color light yellowish olive or grayish olive.
All over upper parts are fine light grayish vinaceous or light vinaceous fawn
dots and on the muscular part of tail but not on the crests. These dots become
larger on dark upper eyelid and give a conspicuous arrangement. In half-
grown tadpoles a cream-colored ring under the eye and extending backward
on either side of back for a distance. In very young tadpoles the ring is almost
complete about the eye but not on the back. This ring faintly present in
mature tadpoles and extends backward on to the base of the musculature of
the tail. Under the eye it is warm buff, on body buffy brown and cartridge
buff or tilleul buff.

Tail. Beginning at body there runs along musculature for one-half or
more of its length a white, cartridge buff or pale cinnamon pink stripe marked
off by the dark brown or black ground color of the upper half of the muscula-
ture and lower crests with prominent large spots black in younger specimens,
and clusters of mouse gray on mature tadpoles. The tip of tail (both upper
and lower crests) clear of spots. Area of crest next to musculature clear in
upper.

General appearance. Tadpole small (36 mm.) full and deep bodied. Tail
medium to fairly long, tip acuminate and very pointed. Tail with a promi-
nent flagellum the dorsal and ventral crests of which are usually colorless.
Tail deep and dorsal and ventral crests well developed. The dorsal crest may
exceed the musculature in depth and extends on to the body to the vertical of
of the spiracle or half way between spiracle and eye. Spiracle sinistral,
directed more backwards than upwards, far below the lateral axis and visible
as an elliptical opening. Eye on the lateral axis, in dorsal aspect on the lateral
outline and in consequence visible from the venter. Anus dextral, very near
the level of the edge of the lower crest. Muciferous crypts indistinct.

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
upper fringe for about 4/1 1 to 2/5 of the length of the upper fringe. The end
of the second row usually is even with the end of the upper fringe. The horny

Hyla femomlis 289

beak is contained about 2.0 times in the upper fringe. The median space be-
tween the lateral second upper labial very short, 6-10 times the length of
either lateral row. The inner papillae extend under the third row of lower
labial teeth making at least two rows of papillae across the lower labial border.
In the lower labial corner there is a heavy papillary series of 4 or 5 rows like
H. versicolor or H. squirella. The lower third labial is long and is usually
about 1. 1 0-1.2 in the first or second row, longer than in H. squirella and is
larger than the horny beak. The first and second rows are about equal and
1. 4-1. 6 greater than the horny beak. This species, like Hyla versicolor and
Hyla squirella has a very angulate upper fringe at its middle.

Measurements. Length of body (8.0-12.0 mm.), in tail (13.5-2.40 mm.)
1.3-2.5, average 1.75. Width (4.5-6.5 mm.) of body in its own length 1.6-2. i,
average 1.8. Depth (4.5-7.0 mm.) of body usually slightly greater than body
width (4.5-6.5 mm.). Depth of body 1.33-2.2 in body length, average 1.68.
Depth (4.5-10 mm.) of tail in length of tail 1.6-2.75, average 2.25. Muscular
part (3.5-4.5 mm.) 1.8-2.3 in depth of tail, average 2.1. Spiracle 1.1-1.4
nearer base of hind legs or vent region (5.0-6.0 mm.) than the tip of the snout
(5.5-8.0 mm.), 1.24 average. Spiracle 1.5-2.5 nearer eye (2.0-4.0 mm.) than
base of hind legs or vent (5.0-6.0 mm.), average 1.87. Eye 1.0-1.75 nearer to
spiracle (2.0-4.0 mm.) than to tip of snout (3.2-4.5 mm.), average 1.25.
Nostril 1.2-2.3 nearer eye (1.2-2.5 mm.) than snout (2.0-3.0 mm.), average
1.85. Mouth (2.0-3.5 mm.) equal to the internasal space (2.0-3.5 mm.).
Mouth contained 1.4-2.0 (average 1.65) in interorbital distance (3.5-6.0 mm.).
Internasal space (2.0-3.5 nim.) contained in interorbital space 1.3-2.0,
average 1.7.

The dimensions of the largest tadpole are:

mm.

mm

Total length

330

Spiracle to vent

6.0

Body length

10

5

Spiracle to eye

4.0

Body depth

5

0

Eye to snout

3-2

Body width

6

5

Eye to nostril

1-5

Tail length

22

5

Nostril to snout

3-0

Tail depth

7

5

Mouth

30

Musculature

of tail

3

5

Interorbital distance

5-0

Spiracle to snout

8

0

Internasal distance

3-5

General remarks. In 191 2 we provisionally identified these tadpoles as of
Acris gryllus because of similar pattern on the tadpole tail and Acris rear
femur coloration. It was not until June 16,1921, almost a month after first
eggs recorded we secured checks to identify these tadpoles. It was not until
July 8 ,we dared describe the coloration provisionally. On July 17, 192 1, we
recorded small tadpoles and only this size. In some ponds July 24, 192 1, the
tadpoles were almost a pure culture of Hylafemoralis and small. On July 26,
192 1, we finally decided the identification and described the coloration of
mature tadpoles.

In 1922, June 26 "in an open pond were a series of Hylafemoralis tadpoles.
They have bright red in the tails of many of them, and long light Hne on the

290 Frogs of the Okefinokee Swamp

middle of the muscular part of the tail. Associates are H. squirella, H.
gratiosa and a few R. sphenocephala. Their tadpoles are smaller than those of
Hyla versicolor."

It might not be amiss to append the description of this tadpole we made
in 19 1 2 then believing it to be Acris gryllus. The description follows but
should not be taken in lieu of the better one which precedes. It is:

"Length of body contained 1.5 to 2.3 times in the tail, average 1.8. Width
of the body in its own length 1.4 to 2.05, average 1.8. Nostrils 1.25 to 2
nearer the eye than snout, average 1.6. Eye lateral, usually nearer the snout
than the spiracle, occasionally equidistant. Distance between external nares
in interorbital distance 1.4 to 2.5, average 1.67; in mouth, .9-1.8, average 1.35.
Spiracle sinistral, usually 1.2 to 1.8 nearer the base of the hind legs than the
snout, rarely equidistant from the two, average 1.37. Anus dextral. Depth
of the tail in its own length 2.0 to 3.7, average 2.9. Depth of the muscular
part of the tail in the depth of the tail 1.5 to 2.8, average 2.17. Mouth is
contained in interorbital distance i.o to 2.0, usual range i.i to 1.6, average
1.25. Greatest length 35 mm. Greatest length of body 11.5 mm. Greatest
length of tail 23.50 mm. Greatest depth of tail 9 mm.

General color of the back (in spirits) yellowish white thickly covered with
fine dark brownish specks or spots or blotches. These spots often appear like
intricate star-like clusters connected with each other at their tips. Entire
venter is yellowish white, the intestine showing through the skin, the general
type of belly coloration reminding one of Hijla versicolor. In the tree toad the
intestine does not show through or is but slightly visible. The dorsal colora-
tion leaves a yellowish white or cream circle about the eye. The distinctive
coloration of this form is that of the tail. The muscular part of the tail has
four distinct longitudinal bands. The first band of deep Vandyke brown or
umber reaches from the body to the tail tip along the ventral edge of the
muscular portion. Above it comes a cream or yellowish white band which
reaches two thirds to three quarters of the distance to the tip. This is followed
by another brown band to the tail's tip. The two brown bands at the caudal
third or fourth of the muscular portion unite and cover this part. Above the
upper brown band is a second short cream band which sometimes runs onto
the body for a short distance. Then the dorsal edge of the muscular part is
more or less brown. The crests are translucent and more or less covered with
aggregate brownish or blackish masses of color. These blotches are more
prominent or intense around the edges of the crests Hke the condition in Hyla
versicolor. These blotches become more numerous as the hind legs develop.
The largest tadpoles have a coloration pattern very much like the above tree
toad tadpoles but without the scarlet or vermilion while the smaller ones
remind one somewhat of the caudal crest coloration of Hyla crucijer. Of
course the muscular bands separate the cricket frog tadpoles at once.

LARVAL PERIOD

In 191 2 we first entered the swamp May 27 or 28 and recorded first activity
at once. Collected spent females May 29 and on July 2 found transformed

Hyla femoralis 291

frogs, or 33-35 days afterwards. This, however, is insufficient evidence. In
192 1, the first spent females were taken April 25, the first positive ovulation
began May 21 and first transformations came June 16, June 21, July 24 and
July 26. This would give us roughly 52, 57, 90, 92 days from first spent female
to transformation dates (June 16- July 26) or 26, 31, 64,66 days from first
positive eggs to transformation dates. In 1922 the first spent female taken
was July 3 and first gravid females June 29, our latest transformations
August 4, 9, 13. These ranges give us 36, 41, 45, 32, 35, 41 days. Possibly all
these transformations are from eggs before June 29. We doubt if the larval
period be shorter than 30 days, and 90 days seems too long. Doubtless 40-70
days are not far from the range.

TRANSFORMATION

Period. In 192 1 our first records of transformation came June 16, July 23,
July 24 and July 26. In 1922 we have transformations June 16, June 21,
June 26, June 27, June 30, July 11, July 12, July 19, July 26, July 27,
August 4, August 9 and August 13. We have outlined actual transformations
from June 16- August 13. The eggs laid August 17, 1922, must transform as
late as October.

Size. On July 2, 19 12 we took on Camp Island a specimen which measured
13 mm. On July 24, 1921, on Billy's Island, in Indian Mound Ponds we took
nineteen individuals 11. 5-1 5 mm. in size. On July 26 we secured three more
13-14 mm. in size. In 1922 we secured two specimens at Callahan, Fla., June
16, 1922, 14 mm. and 15 mm. In the Okefinokee Swamp in 1922 we secured
31 transformed individuals, 10 mm. -14 mm. in size. In all we measured 57
individuals with a range of size 10-15 rnm- The major mode was 13 mm. (15
specimens) and two secondary modes of 12 mm. (12 specimens) and 14 mm.
(12 specimens), and the average of the 57 specimens was 13 mm.

In 1928 we discovered several tadpoles labelled Acris which are really Hyla
femoralis. One lot, June 16, 1928 is a transforming series.

m. 4 legs. Tadpole mouth.
' " " Teeth rows gone.
" " Tadpole mouth
half changed.
" ii.o " " 14.5 " " " Tadpole mouth no

teeth.

On July 23, 192 1 we find tadpoles and 4 transformed frogs and beyond, one
II. 5 mm., 12 mm., 13 mm., and 14 mm.

General remarks. At transformation many of the tree toad species tend to
be green with none of their distinctive markings. In the first identification of
these forms the tails of the series not fully transformed often helped materially.
Adults of Hyla femoralis are not often green (sometimes so) but their trans-
formed life frequently starts with a green livery. They are then hard to
separate from Hyla squirella, Hijla cinerea, etc. On July 18, 1922, "around
the edge of a sphagnum bog overgrown with bushes and trees we found trans-

3d3

J II

. 5 mm.

Tail 14

>)

II

•5 "

" 12

>>

II

.5 "

" 9

292 Frogs of the Okefinokee Swamp

formed Hylafemoralis. They were green above." Or on July 27, 1922, "in a
pond where we heard Hyla gratiosa and took transformed green Hyla gratiosa
Mr. Pirnie found two transformed Hyla femoraUs, golden green and resting
on stems 1-3 inches above the water. If they were shaken off they swam to
another stem and climbed up. They were a prominent green but when trans-
ferred to a cocoa can they changed in five minutes to an almost black color."

GROWTH

In 191 2 we secured the following material: "On May 28-29, 1912, we took
one specimen, 17.75 mm. long, another on June 6-7, 19 12, 21.25 mm., a third
June 24, 19 1 2, 21 mm., a fourth from the edge of the swamp Sept 11, 1909,
18 mm., and a fifth, 16.5 mm. These measurements 16. 5-2 1.2 5 mm. from
June i-September 11 represent 10-12 months' old frogs." On February 9,
191 1, Professor J. C. Bradley took a 20 mm. at Waynesville, Georgia.

In 19 1 2 on June 6-7 when a 21 mm. was collected two more were taken
31 and 36 mm. respectively. On June 24 besides the 21 mm. specimen we
have a 27 mm. and a 31 mm. specimen. On June 5 when a 25 mm. was taken
we also recorded 33 and 36 mm. specimens. On June 15 two specimens we
have respectively 26 and 31 mm. Possibly on this basis the one year olds
extend from 16.5-26 mm.

In 1921 we captured transformation sizes from 11. 5-15 mm. On April 25
when we took a 20 mm. specimen we also captured 24, 26, 26, 28, 30 and 33
mm. specimens. On June 3-5 we had a series 20, 24, 25, 27, 27, 27, 30, 30, 31,
31, 32, 33, 33, 34, 34, 37, 37, 37, 4o mm. respectively. Or on May 21 a series
of four yielded 25, 28, 31, 33 mm. specimens. With the other material col-
lected in 192 1 the line between i and 2 years seems to be from 24-26 mm.,
while :n 191 2 it is apparently 26-27 mm.; the line between second and third
years in 191 2 between 32-34 mm., and in 192 1 between 32 and 35 mm.

In 1922 we took one specimen of 23 mm., two of 29 mm., and one of 34 mm.
On August 13 we secured a 19 mm. specimen on the Sugar House, St. Mary's,
Georgia. All the other lots begin with 32 or 33 or 29, 29, 28, or 27 mm. Two
series are: July 21-27, 28, 28, 28, 29, 30, 30, 30, 30, 31, 31, 31, 32, 32, 32, 32,
33, 33, 34, 34, mm.; July 11, 28, 28, 29, 30, 30, 31, 38 mm. The 1922 material
apparently separate at 25-26 mm. for the line between ist and 2nd years and
at 33-35 mm. for the Une between 2nd and 3rd years.

Considering seasonal, yearly or sexual variations the evidence points to
10-15 mm. for transformation size, 17-26 mm. for ist year olds; 26-32 or 34
for 2 year olds; 32 or 34-40 mm. for 3 year olds.

FOOD
This phase of its natural history we have not examined.

ENEMIES
In 191 2 we summarized it thus: "This species was represented in the
stomach contents of one black snake {Coluber constrictor)oi onb southern
riband snake (Thamnpohis saimtus sackeni), of two garter snakes {T. sirtalis)

Hyla femoralis 293

and of one pilot snake {Elaphe o. confinis). In other words, two distinctly
terrestrial species ate it, one aquatic species and one more or less arboreal form.
The last, the pilot snake may have captured it in the piney trees, but all the
rest doubtless took it on its journey to the breeding pools or while in them,
or this species may be more terrestrial in other months and not exclusively
arboreal as previously held. This tree frog may be more frequent in the
reptilian diet of the swamp because it is more common than its relative, the
Carohna tree frog (Hyla cinerea)." On July 2, 1922, we saw at night a small
garter with an adult Hyla femoralis in its mouth. The snake had it head first.
On June 27, 1922, in one pond where we captured the Hyla gratiosa tad-
pole with bifurcated tail and Hyla squirella tadpole with one hind leg we found
a Hyla feinoralis tadpole with only one hind leg. Another Hyla femoralis tad-
pole had the forelegs about to burst out and no hind legs at all. Was it ab-
normal, pathological or were they snapped off by turtles (Deirochelys reticu-
laria) or snakes which we captured in this pond?

AUTUMNAL DISAPPEARANCE

The latest records we have are for mid-September, 1922, and August 22,
192 1. Some would have transformed in October. Doubtless this month or
November are the periods of beginning of inactivity.

AFFINITIES

Holbrook (1842, Vol. IV, p. 128) alludes to Dumeril & Bibron's mistake
as follows: "Dumeril and Bibron consider this animal as identical with the
Hyla squirella, from which it is, however, perfectly distinct.

1. It is about two-thirds the size (as a matter of fact H. femoralis
is larger, A. H. W.).

2. Its general colour and markings are different: there are no white
lines along the upper lip, and the yellow spots on the thigh always exist
there, and never in the Hyla femoralis.

3. It differs in habits, for it is never found near outhouses or about
fences and in old fields."

In 185s John LeConte (1855, p. 428) refers to the same matter. "It is
wrong in Dumeril & Bibron to say that this species is a variety of Hyla
squirella. In shape and size the difference is not considerable. The latter
animal during the warm season is always to be met with about the houses, the
H. femoralis never. Besides, their notes are entirely different." As late as
1882 Dumeril & Bibron's mistake somewhat influences Boulenger (1882, p.
398) in his statement "Appears to be specifically distinct from H. squirella.^'

This species seems in adult appearance to be a small species of the Hyla
versicolor, yet is readily distinguishable. The individual eggs look Hke those
of this species and the surface films on the water appear to be of the same sort,
though Hyla femoralis may lay larger films. Of the tadpoles we have the
note (June 27, 1922) that "The tadpoles have the same bright red color in

294 Frogs of the Okefinokee Swamp

tail as Hyla versicolor. Are they related? The color pattern of the tail in
alternate light and dark, longitudinal bands is like that of Hyla andersonii
and Pseudacris species. In mouth parts the tadpole falls into the group of
Hylas with long hind lower labial with rows like Hyla versicolor, H. squirella
and Hyla arenicolor.

BIBLIOGRAPHY
1907 Brimley, C. S. Journ. Elisha Mitchell Soc, Dec. 1907, Vol. XXIII, No. 4. pp.

158, 159-
1889 Cope, E. D. U. S. Nat. Mus. Bull. No. 34, 1889, pp. 371, 372.

1914 Deckert, R. F. Copeia, Feb. 14, 1914, No. 3, p. 3.

1915 . Copeia, May 15, 1915, No. 18, p. 3.

1862 DeKay, J. E. Zoology of New York, Part III, p. 72.

1906 Dickerson, M. C. The Frog Book, pp. 150, 151.

1842 Holbrook, J. E. North American Herpetology, 1842, Vol. IV, pp. 127, 128.

1856 LeConte, John. Proc. Acad. Nat. Sci. Phila., Dec. 1855, pp. 428, 429.

1924 Myers, George S. Copeia, June 30, 1924, No. 131, p. 60.

1924 Schmidt, K. P. Copeia, July 15, 1924, No. 132, p. 68.

1923 Stejneger, L. and T. Barbour. A Check List of North American Amphibians and

Reptiles. 2nd Edition, Cambridge, 1923, p. 30.

1923 Viosca, Percy, Jr. Copeia, Feb. i, 1923, No. 115, p. 36.

1920 Wright, A. H. U. S. Com. Fish. Report 1919, Doc. 888, p. 29.

1923 Wright, A. H. Copeia, Feb. i, 1923, No. 115, p. 34.

1926 Wright, A. H. Ecology, Jan. 1926, Vol. VII, No. i, p. 82.

1923 and A. A. Wright. The Anatomical Record, Jan. 20, 1923, Vol. 24, No. 6,

p. 406.

1924 Wright, A. H. and A. A. Wright. The American Naturalist, July-August, 1924,
Vol. LVIII, p. 378.

Hyla gratiosa LeConte

(PI. II, Fig. 4; V, Fig. 3; VII, Fig. 6; X, Fig. 5; XII, Fig. 2; XV, Figs. 4, 13-16; XVII;
XXIX; XXX; Text Fig. 11)

COMMON NAMES

"The Barker." "Barking Frog." "Coat Bet." Florida Tree Frog. Georgia
Tree Frog. Florida Hyla. Florida Tree Toad.

RANGE

Check list. "Type locality: 'Lower plantation of Georgia,' evidently one
of the LeConte plantations in Lloyd or Liberty County. "Range: South
CaroUna to Florida and Mississippi." — Stejneger and Barbour Check List,
1923, p. 31.

Supplementary records. In L.ousiana Viosca (1923, p. 37) found this
species in the "Longleaf Pine Flats. . . . Typically the country is north Flori-
dian rather than Louisianian, and several southeastern species, not found else-
where in our state, have congenial habits here. Bufo quercicus, Bufo terrastris
and Hyla gratiosa are significant examples." Brimley, C. S. (1907, p. 158) is
quite justified in including this species in his Key to the Species of Frogs and
Toads Liable to Occur in North Carohna."

The Wilmington type of country very likely will yield it some day. Seven
miles north of Millen, Jenkins Co., Ga., Mr. M. D. Pirnie and I started "for
an immense chorus of Hyla gratiosa. We travelled about 1/2 mile and the
chorus seemed 1/2 mile farther on. It was in a dense tangle of a swampy

Plate XXIX

Florida tree-frog ( Hyla gmtiosa)
Cornpatch, Billy Id., Ga. June 6. 192 1.
Frog on sand, Billy Id., Ga. June 5, 192 1.
3. 4. 5. Males croaking in overflowed cornfield, Billy Id., Ga. July 3, 1921.
6. Mature tadpole with forked (abnormal) tail. Petty pond Trail Ridge, Folkston,
Ga. July 31, 1922. Lateral aspect, x 1.3.
Mature tadpole, (normal) Chesser Id., Ga. July 3, 1922. Lateral aspect, x 1.3.

Hyla gratiosa 295

stream." Just north of Screven, Wayne Co., Ga., the evening of June 11,
192 1, we heard Hyla gratiosa. In 1906 Miss Dickerson (pp. 124, 125) writes
"unfortunately this tree frog has an unusually limited range. It is very
rarely found outside of Florida." The last twenty years has extended its
range considerably.

Local Okefinokee records. In 1 9 1 2 we wrote of this species as follows : "This
species was on hypothetical list and escaped us from May 30- July 15, 191 2.
Between July 15-Nov. 15, 1912, the Lees secured one fine male of the Florida
Tree Frog. LeConte (Cope '89, p. 379) had this from Riceborough, Ga., and
C. B. Adams (Cope '89, p. 379) took it in Georgia while Brimley's ('10, p. 11)
records came from Florida and Mississippi. The present record being inter-
mediate helps to bridge the gap between these two areas."

In 192 1 we captured or heard it at Billy's Island, Chesser Island, Fargo,
south of Moniac in Florida, near Macclenny, Florida. In 1922 we recorded
it on Chesser's Island, near Starling Branch, on road from Chesser Island to
Folkston, along road Folkston to Moniac, and in the country generally from
swamp to St. Mary's River. It is scarce within the swamp, more common on
its outskirts outside the swamp.

GENERAL APPEARANCE

Brimley (1907, pp. 157, 158) characterized it as a large tree frog with no
X marks on back, no lateral stripe of yellow, no vermiculations, spottings,
variegations of yellow on rear of thighs, but with the feet edged with j^ellow.
Miss Dickerson (1906, p. 125) holds " Hyla gratiosa presents a pecuHar appear-
ance because of the even granulation of the upper parts. In this characteristic
of the skin it differs from all other North American Hylas. All others present
a surface either smooth and fine in texture, as in the case of the Green Tree
Frog ( Hyla cinerea) or rough with fine warts, as in the Common Tree Toad
{Hyla versicolor)."

COLORATION OF SPIRIT SPECIMENS (1912)
The coloration of our first specimen of 1 9 1 2 : In spirits purplish brown on
the back with prominent darker elliptical or round spots. These may be solid
or with lighter centers or with a smaller yellowish spot on them. Tibia im-
perfectly barred while the bars of the thighs are on the anterior half of the
dorsal surface, the posterior half being plain with no bars or spots. Forelimbs
with two or three small dark spots on interior edges; fore-arm with two yellow-
ish or whitish bands along its posterior edge; upper labial stripe yellowish
or white. The specimen we have is a male and has the prominent vocal sac
area of the same color as the back, the breast fold being yellowish as on the
rest of the under parts.

STRUCTURAL CHARACTERS

(Based on our lone specimen of 191 2, we wrote these notes.)
Upper parts except arms and legs strongly granulate; under surface of
hindlimbs only granulate on posterior half of thigh, otherwise smooth; under
surface of arm, prominent breast fold and lower throat smoothish or slightly

296 Frogs of the Okefinokee Swamp

granulate; chin granulate; webbing of hind feet to the discs which are very-
large, 3 mm., or .5 of the tympanum; discs of fore feet, 4 mm., or .66 of the
tympanum; a dorsal fold over the ear to the shoulder and continued often as a
loose fold along the side of the body half way to the groin; a tarsal fold;
vomerine teeth between inner nares; head ahead of eye short, obtuse; vocal
sac well developed.

MEASUREMENTS

(Recent Material)

Head to angle of mouth 1.16 (20 mm. transformation) — 1.15 (28 mm.) —
1.35 (44 mm.) — 1.3 (56 mm.cT) — 1.3 (68 mm. cf) in width of head; head to
rear of tympanum (20 mm. lacking) — i. 4-1. 05 — 1.15 (56 mm. cf) — 1.08
(68 mm. cf) in width of head; head to angle of mouth 3.33-2.8-3.14-3.3-3.4
in length of body; head to rear of tympanum ( )-2. 54-2. 4-2. 9-2. 8 in length
of body; snout .6-.8-.833-1.0-.91 in first finger; snout . 8-1. 0-1.2 7-1. 11-.9 in
fourth finger; snout .5-.6-.72-.84-1. in first toe; eye i. 25-1. 25-1. 5-1. 5-1. 87
in snout; eye ( )-.5-.66-.83-i.o7 in tympanum; eye .75-1.0-1. 25-1. 5-1.0
in first finger; tympanum ( )-2. 50-2. 25-1. 8-1. 7 in snout; tympanum
( )-4. 75-3. 75-2. 4-3.0 in intertympanic width; internasal width i. 25-1. 2-
.8-1.0-.91 in upper eyelid width; interorbital width .71-.85-.51-.77-.66 in
upper eyehd width; interorbital width .57-- 71-- 7 7-. 7 7- 73 in internasal width;
interorbital width ( )-2. 7-2. 3-1. 84-2. 6 in intertympanic width.

Forelimb: Forelimb 1.53- 1.7 5- 1.8 2-2. 0-2.0 in length of body; forelimb
2.0-2.75-2.66-2.6-2.73 in hind limb; first finger i. 66-1. 62-1. 66-1. 33-1. 25 in
third finger; second finger i. 25-1. 44-1. 37-1. 2-1. 35 in third finger; second
finger .75-.88-.83-.9-1.09 in first finger; third finger .8-.92-.84-7. 5-8 in sec-
ond toe; fourth finger i. 25-1. 3-1. 08-1. 2-1. 36 in third finger; fourth finger
.625-.6-.56-.75-.72 in first toe; internasal width i. 5-1. 6-1. 5-1. 8-2. 18 in
first finger; internasal width 2.0-1. 8-1. 8-2. 0-2.0 in second finger; internasal
width 2.5-2.6-2.5-2.4-2.72 in third finger; internasal width 2.0-2.0-2.3-2.0-
2.0 in fourth finger.

Hindlimh: Length i. 3-1. 57-1. 45-1. 3-1. 36 in hindlimb; tibia i. 9-1. 64-
1.9-2. 19-2. 12 in length; tibia 2.47-2.6-2.78-2.86-2.9 in hindlimb; tibia 1.23-
.94-1.04-1.09-1.06 in forelimb; tibia .81 .73-.87-.98-.93 in hind foot; first
toe 1. 6-2.0-1. 6-1. 2-1. 5 in second toe; 2.6-3.0-2.3-1.86-2.37 in third toe; first
toe 3.2-3.06-3.0-2.0-1.9 in fourth toe; first toe 2. 4-3. 3-2. i-i. 6-2.0 in fifth
toe; second toe i. 62-1. 5-1. 45-1. 55-1. 58 in third toe; second toe 2.0-1.8-1.86-
2.0-1.9 in fourth toe; second toe i. 5-1. 66-1. 34-1. 33-1. 25 in fifth toe; third
toe 1. 23-1. 2-1. 36-1. 3-1. 2 in fourth toe; third toe .92-1.1-.93-.85-.84 in fifth
toe; fourth toe .75-1.13-1.05-.66-.69 in hind foot; fourth toe 1.31-1.54-1.21-
1.41-1.39 in tibia; fourth toe i. 62-1. 45-1. 26-1. 55-1. 5 in forelimb; fifth toe
1. 33-1. i-i. 35-1. 5-1. 43 in fourth toe; internasal width i. 25-1. 2-1. 3-1. 5-1. 45
in first toe; internasal width 2.0-2.4-2.1-1,8-2.18 in second toe; internasal
width 3.25-3.6-3.0-2.8-2.34 in third toe; internasal width 4.0-4.4-3.8-3.6-4.1
in fourth toe; internasal width 3.0-4.0-2.8-2.4-2.9 in fifth toe.

Hyla gratiosa 297

HABITAT

In 1856 John LeConte (1856, p. 146) in his description writes of his three
specimens as follows: "One of them was taken in the water of a pine barren
pond, another was found in a cavity of a sand pit, and the third upon a tree in
the forest." In 1922 H. P. Loding (p. 19) makes an interesting note about its
great abundance. "Thus, the largest of our tree frogs seemed to be very rare
previous to the hurricane of July, 1916, up to which time in all our collecting
only two adults and two very young specimens were taken; but suddenly
during August of that year the species became very common, in fact could be
seen by the thousands in the Satsuma orange orchards throughout Mobile
County. Since then it has gradually become scarcer and at present is not seen
very often."

We do not know whether its non-ovulation habitat be primarily the pine
woods, the hammocks, the fringe of cypress ponds on the edge of cypress
bays. In each habitat we have records of "barking frogs."

FIRST APPEARANCE

In 1856, John LeConte, (p. 146) writes: "During the last spring, whilst I
was residing in the lower country of Georgia, it was my good fortune to meet
with three specimens of the animal described below." Deckert (1915, p. 5)
records them in March. T. Van Hyning (1923, p. 68) writes that "On the
night of March 18, while some of the boys of the biological class of the Univer-
sity of Florida, Gainesville, Fla., were collecting frogs, among other species
taken, were thirty specimens of the Florida Tree Frog Hijla gratiosa LeConte ..."

GENERAL HABITS

Deckert (19 15, p. 4) writes that ^' Hyla gratiosa LeConte, the Florida tree
toad, is a handsome species, and the largest of the North American tree toads,
reaching a length of 3I inches from snout to vent. It is heavily built, with
large adhesive disks on fingers and toes. It differs from all other species in its
evenly granulated skin, and regular pattern of roundish spots. The color
is ashen-gray, purplish or green of some shade. The spots are darker than the
ground color, evenly distributed over the open surfaces, and may be absent
when the frog changes to pale golden green. The arms and legs are banded.
The throat of the male is rich chrome yellow or green."

Dickerson (1906, pp. 125, 126) studied this frog carefully in captivity.
She observes that: "The Florida Tree Frog has colour changes as marvellous
as those possessed by most of the Hylas. It may be plain bright green and re-
main so for months, in fact, one in this phase of colouration was kept for over
two years and during this time there was no tendency to become spotted or
to turn brown. Also, the brown-spotted phase may likewise endure for months
without change. On the other hand, the changes may be very rapid indeed.
The tendency for rapid colour change, here as elsewhere among the Hylidae,
seems to be at its height when the animals are well-fed, and in the season of
greatest activity, that is, in the spring and summer months. At this time the

298 Frogs of the Okefinokee Swamp

greatest variation in colour may be seen in an individual during the twenty-
four hours. It may be bright green with vivid spots, dark at their edges and
light in their centres. Gradually (within the space of ten minutes) the spots
become more and more obscure until the frog is plain green, very light in tone.
Again, within the half-hour the spots reappear, become more distinct, darken-
ing in their centres. The ground color becomes ashy in the middle of the back;
this colour darkens into a mulberry, which colour spreads, obscuring the
green. Meanwhile the spots continue to darken until the whole frog is rich
brown prominently marked with darker brown spots."

"The throat-pouch of the male is very large and is bright yellow or vivid
green in colour. When the tree frog is taken suddenly into the hand, this sac
is likely to be inflated and the frog gives a harsh-sounding squawk. The voice
is usually harsh and low-pitched."

"This Hyla is a curiously artificial-looking frog. If it did not move, we
might well think some one's fancy had moulded it out of wax. When it is
angry or startled, it fills the lungs, expanding the body until it is nearly as
broad as long. It has the slow tendencies of some of the other large tree frogs.
It shows a marked desire to cling to one's finger and will not leave, even when
the finger is constantly turned so that the tree frog is head down. It will each
time clumsily turn around to bring its head uppermost, tuck its toes well
under, and settle for a period of contentment and rest."

We observed that the first male which we captured croaking in a corn field
June 5, 1927, would squawk. On June 7, 1927, we noted that "the male would
open his mouth once in a while after I captured him. He was in a glass can in
my pocket and several times he spoke thus. Later during the next day it
would thus do if taken by the legs and pressed ahead of hind legs. Thus held
while rubbed on belly or back it will swell up so that it looks perfectly round
from above. Often it has a curved appearance; it will bow its head down to
ground and back be arched. It can leap i-i 1/2 feet, possibly 2 feet in
awkward, cumbersome fashion. It has a decided secretion from the skin."
On July 16, 192 1, one of the Chesser boys noticed its odor. He pronounced
it bad and "tried three different times to get it off his hands."

VOICE

Deckert (1915, pp. 4, s) considers "It is not very plentiful anywhere, and
rarely met with outside the breeding season. During the latter time it comes
down out of the trees and small companies of from four to ten specimens, in
widely scattered pools or "bayous" attend to their breeding duties amid the
"loud calls of the males. The call can be heard for over a mile, and sounds
like a large gong, or church bell, being of unusual depth, and very clear, with a
second's interval between each dual note. The male while singing, floats on
the water, the large vocal sac throwing the frog into a vertical position with
every utterance."

The first time we heard this species to know it as such was on June 5, 192 1.
After a period of heavy rainfall "We visited in the evening an oak-toad pool

Hyla graiiosa 299

near the cultivated fields of Billy's Island, and heard a curious ionk, tonk, like
some one pounding on a hollow, heavy barrel or hogshead. The call was
woody, deep. At first it reminded me more of a frog (Rana) than a tree frog
(Hyla). It seemed at first somewhat like the call of Rana clamitans only too
many times separated and actually the resemblance is only fleeting and shght.
Later we found other croakers. One was on the raised ridge of land beside a
corn stalk. It was green with the spots usually figured. Another cf^ was
lying flat in the water at the edge of a weedy ridge. It was absolutely flat and
spread out. It was without the spots and uniform duU, brownish-green like
the color of the water. Another was in a furrow between two rows of corn.
It was spotted and alert. First one rested on ground more or less horizontally;
the second in water horizontally, and the third more or less diagonally upright.
The call can be heard at a long distance." Usually the heels are widely
separated at croaking and the lower part of the body dips somewhat.

On July 3, 1 92 1, on Billy's island "after a great downpour of rain at 8:00
p.m. heard a cf Hyla graiiosa calling over in the Hammock. Decided to go
out. Found another male near camp in temporary pool. No end of Hyla
femorah's, Hyla cinerea, Gastrophryne, Bufo terrestris and Chorophulis ocularis
calling. Found a Hyla gratiosa cafling but I went past it. When I returned it
had moved 10-15 feet away from its first position. It was of a greenish cast
of olive in color.

"Found a male H. gratiosa beside a grassy bank. He was as big and round
as the top of a Mason jar. How his throat would puff out. Was an easy
subject to move into good position. Used either a big stick or flashlight or
other means."

F. Harper makes these journal notes on this subject. "56 calls in a minute
quite regularly spaced. Then 59 to a minute not absolutely regularly spaced.
Then 55 to a minute. Even between calls, keeps body puffed out. Inflation
of body and throat in calling, alternate as in Hyla cinerea. There is a perceptible
pause between each note in cafling, the throat meanwhile remaining at one
quarter its full inflation. The inflation and deflation are accomplished very
rapidly, making it quite a trick to catch a good inflation with the flashlight.
The throat remains at about this quarter inflation during a considerable
pause in calUng. Very brownish at night."

On July 15, 192 1, on Chesser Island we heard of "Coat Bet" frogs. The
first day we thought they must be Rana virgatipes but the following evening
we found them to be Hyla gratiosa. The people on Billy's Island have no
common name for them. On Chesser Island they are called Coat Bet be-
cause that is the way it sounded to the Chessers. They said the "Coat Bet"
caU was produced by one frog calling right after another.

On July 3, 1922, in a temporary pool "1/2 mile west of Trader's Hill, Ga.,
heard 4 or 5 Hyla gratiosa. Some would stop calling when we approached
and if we came too near they ducked under water and swam away. Finally
we captured two. One looked green and one brown."

In conversation with Mr. P. A. Chesser he spoke of "Coat Bets" as follows
(July 15, 192 1). "Sometimes in rainy weather we hear these around the

300 Frogs of the Okefinokee Swamp

house and on the Island. Real season is summer time when there is plenty of
rainy weather. 'Spanish Curlews' once fed on them."

While we were in the swamp we usually recorded this species either as
Barker, the Barking Frog or Coat Bets. The last refers to the normal note in
the breeding pools; the first two to a puzzle that perplexed us for two seasons.

On May 22, 192 1, we ''heard in a tree a curious note or croak in the ham-
mock near Newt Pond. After a time it croaked more loudly from a moss-
laden black gum and was different. At first I thought it an aberrant H.
cinerea. Is it Hyla gratiosa?" June 5, 1922, in a cleared field near this
hammock we found 2 or 3 male Hyla gratiosa calling in regular fashion. On
July 16, "during the morning we heard a barking frog in the trees south of
camp. We are almost satisfied tonight that it is Hyla gratiosa." That night,
in a nearby pond was an immense chorus of "Coat Bets" (Hyla gratiosa).

In 1922 the barkers perplexed us even more. Mr. R. A. Chesser told us
"the Barking Frog some folks say is a red-headed scorpion. I believe it is a
frog." On June 24, 1922, at the old railroad pools just south of Trader's Hill
"heard this form. It is in a grove of trees. One of our party wonders if it is
Hyla andersonii?" On June 26, the writer heard one in the high long-leafed
pines north of Starling Branch. On June 27, "at Camp Pinckney heard a
barking like frog. Later, on wild rice in the direction from which the noise
came was a curiously colored Hyla cinerea. Is it the barker? Does it call
differently when in high trees and when in breeding waters?" On July 2
"heard a barker near Anna's Pond." On July 11 and July 20 they were
barking on Chesser Island. Not until July 26 did we solve the puzzle. About
three miles along on the road from Chesser Island to Folkston we heard in the
evening "in a cypress pond to the right of the road some Hijla gratiosa and
beyond them, a barker or two. Went after the barker. Found one in a small
gum 4-5 feet, possibly 6 feet up. It is Hyla gratiosa! I saw him do it. Two
more barkers beside one I caught. Several Hyla gratiosa in water calling
normally. Is the barking note the call of the trees before reaching water
level. There is a great chorus of Coat Bets (normal caUing frogs) to the north-
ward." On August 16, 1922, at Hilhard, Florida, where we found both
spadefoots and Florida tree frogs breeding, 2-6 p.m., we heard barkers in
the pines near the place. At 7.15 p.m. Coat Bets began to croak in the trees,
but at times they were "Coat Bets" and at other times "Barkers." "Coat
Bets" and "Barkers" are one, Hyla gratiosa.

A resume of the croaking dates for 192 1 and 1922 are as follows:

1921

May 22. Curious croak or note from moss-laden black gum in hammock,
Billy's Island.
3 croaking, Billy's Island.

cf calling in Hammock. Another in overflow area near camp.
During the morning heard barking frogs in trees south of camp.
Immense chorus of H. gratiosa.

June

5'

July

3-

July

15

July

16.

Hyla gratiosa

301

July

17-

July

29.

Aug.

3-

Aug.

4-

Aug.

6.

Aug.

7-

June

10.

June

II.

June

12.

June

14.

June

24.

June

26.

June

27.

July

2.

July

3-

July

II.

July

12.

July

13-

July

20.

July

25-

July

26.

July

27.

Aug.

8.

Aug.

10.

Aug.

II.

Aug.

13-

Aug.

16.

Aug.

20.

Calling loudly.

Calling on Billy's Island.

Heard at night at Fargo.

Heard at night at Fargo.

Big chorus near Moniac.

More heard south of Moniac in Florida.

1922

North of Millen, Ga., large chorus.

Near Screven, Ga., heard several.

Chesser Island, Ga., no end of Hyla gratiosa calling.

Several calling but not in chorus.

Heard near Trader's Hill in grove of pines.

A barker in a high pine.

Camp Pincknej^ i or 2 barking frogs.

Near Anna's Pond barking frog. Also near Starling Branch.

Near Trader's Hill several in flooded fields.

Heard a barker on Chesser Island.

Spent an hour searching for a barker in pine tree.

Barker calling in hammock.

Calling 2:00 p. m.

Heard i or 2 north end of Island about 9:00. Temperature 72°.

Solved barker. It is H. gratiosa. Heard barking and normal notes

in same pond.

9:00 a. m. Air 84°. Plenty calling.

Along Old Okefinokee Road several heard.

One calling near Chesser School.

None heard.

Very few heard Folkston-Chesser Island, 9-10:30 p. m.

Several calling in pines and ponds, 2-6:00 p. m. Croaking 7:15.

Heard a few on Moniac-St. George-Chesser Island Road.

In 192 1 on May 21 it rained almost continually; on May 22, we heard
our first H. gratiosa on Billy's Island. On June 4 we had a heavy downpour.
The following day we found three croaking in an overflowed corn field. On
July 3 after a week of rain it poured ; that evening four males began croaking.
On July 15, on Chesser Island a thunder storm came in the afternoon. That
evening the Florida tree frogs began a chorus. The following day, July 16,
another storm came and another congress came. On July 27 and 28 we
were flooded and the evening of July 29 the frogs began agan.

Weather stations around the swamp. Rainfalls of 1.22, 1.78, i.oo, 1.47,
2.40, .91 inches respectively usually came before the croaking of this species in
192 1. The air minima are from 66-70°. The maxima are from 78-96°F.
Warm rains are the important factor. In 1922 when croaking records come
the lowest and highest minima are 62 and 83° and the lowest and highest
maxima 81 and 98°. Most minima are from 66-76°, the maxima 87-94°,
Most of the large choruses came after rains of 2.05, 2.00, 1.60, 1.35, 1.45
inches. The average of the minima is 70°, of the maxima 91°. In 1922 as in
192 1 high air temperatures and dampness produced choruses in midsummer.
Of the early spring we cannot speak.

302 Frogs of the Okefinokee Swamp

MATING

Male {From life, June 5, 1921). Lettuce green on hind legs and forelegs,
oil green or cerro green on the back. The encircled spots of the back have a
ring of mars brown or bone brown or other browns with green ground color
within. There are occasional spots as big as pin heads or twice of bright green
yellow or greenish yellow or green yellow. This color is on the groin to just
back of arm insertion back of which for one half of an inch is a clear whitish
area. The greenish yellow from groin forward demarcates side color from
pink white or creamy belly color. From tip of snout along upper jaw under
tympanum and along side to within i inch to 1/2 inch of leg insertion is a
pinkish white stripe. Below it is a line of mars violet or taupe brown (a
purplish brown class of color) which expands behind the angle of the mouth
into a large area on the side. Same color back of pinkish white or white
area back of brachium and antebrachium onto the last finger. Same white
line around vent with mars violet below white line. Same combination on
the knee. Two or three white patches on foot behind with mars violet
between and behind them. Back of femur or thigh dull Indian purple. No
spots. Below the expanded mars violet area of the side is a small whitish
line 1/2-1 inch long, then comes greenish yellow and finally the belly color.
Just back of chin is grayish white speckled with taupe brown. Then comes
lettuce green or oil green from angle of mouth around across to angle of
mouth, the band 1/2 inch wide when not inflated. Then comes wrinkled part
with light orange yellow or deep chrome. This color more or less back to
pectoral fold of skin from forearm to forearm.

Iris spotted purplish black and bronzy or some kind of vinaceous.

Male {From life, July 17, 1921). The Hyla gratiosa males in botany drum.
Upon opening the can one was all green with a few yellowish spots but not the
regular circles, the stripe along the jaw and on the side reminding one of the
same in Hyla cinerea. Hardly any white or purplish shows anywhere. An-
other was a vinaceous gray with the rings. Another dark green and the others
still different.

Female {From life, July 17, 1921). Belly and under side of legs white, a
few picric yellow spots on side below band, also rear end of white line on side
becomes picric yellow in the groin. Picric yellow in the axilla of forelegs and
on either side of light central throat and chin region which is white or sulphur
yellow with a few spots. Pectoral region sulphur yellow. Stripe around snout
along upper jaw to groin white except for rear and not interrupted as in the
male. Spots on back really black encircled. Color of side and chin, spots,
rim of jaw, bands on arms hays maroon, chocolate or warm sepia or bone
brown. Upper parts calliste green or casse green or apple green.

Structural differences. The color difference of the venter is the most
striking difference. One female (68 mm.) long has no hays maroon on the chin
as in the color description of the female (July 17, 192 1). In the mated pair
described the female has the granulations a little less pronounced on the
ventral parts, thighs and dorsum than its mate, but the large female above
has granulations on venter the least pronounced of the 19 adults we have.

Plate XXX

Florida tree-frog ( Hyla gratiosa)
Coat-bet pond, Chesser Id., Ga. July i8, 192 1.

2. Male, Billy Id., Ga. June 6, 1921. Semi-inflation. Lateral aspect, x 0.4.

3. Male, Billy Id., Ga. June 6, 192 1. Before rubbing, x 0.4.

4. Same, after rubbing, x 0.4.

5. Female and male, Chesser Id., Ga. July 2t„ 1922. x 0.4.
Pair on log, Chesser Id., Ga. July 17, 192 1.
Male, Chesser Id., Ga. July 23, 1922. x 0.8.
Egg mass in a pan, Chesser Id., Ga. July 18, 1921.

Hijla gratiosa 303

On the back they are much reduced in prominence. The upper hind legs and
sides have practically none. Our largest female is 68 mm., largest male 68
mm. The smallest male we took is 52 mm. They must show male coloration
much smaller than 52 mm. Doubtless the males and females begin breeding
three years from transformation. After we had started to measure the two
largest frogs each 68 mm. we noted that the male had a tympanal diameter of
7 mm., the female 5 mm. This is the first instance of decided tympanal
difference we have noted in Hylas. In the same specimens the male has the
two outer fingers of the hand thicker and broader than the similar fingers of
the female.

Our first record of mating came the evening of July 16, 1921. "We went
out 8:30-10:00 p.m. south of camp to a pond (Coat Bet Pond). Immense
deafening choruses of Hyla gratiosa, Pseudacris ocularis, Hyla femoralis,
Biijo quercicus and Acris gryllus. We looked for Coat Bets {Hyla gratiosa).
One male was on a cypress limb touching the water; another on a tussock
of sedges; another on water amongst grass. We caught 9 or 10 males. The
pond is generally shallow but water was generally 8 inches-i 1/2 feet deep in
places." Found the eggs next morning. In the evening July 17, 1921,
"Coat Bets were calling loudly and presently on a log next to the water's sur-
face saw a mated pair of Hyla gratiosa. I held my breath but caught them
very easily in my can. Thought I never would get the can out of my pocket.
When I returned to camp the pair had broken and we put another male with
her. One of the Chesser boys presently found a mated pair of Coat Bets on a
log. We set up the camera and they leaped away but swam no more than
three feet and hid under a fern leaf. I caught them and put them back on the
log. Held my hand partially over them while Harper focussed. Then I raised
my hand one foot above them ready to seize if need be. Harper trigged the
flash a dozen or two times before it went off. The pair remained just as before.
When we put them in a can they did not separate."

On August 16, 1922, after a severe rainy period in one of the shallow,
temporary spadefoot pools "were several pairs of Hyla gratiosa."

All our records of mating came at night.

Amplexation. The amplexation of every pair caught was axillary. So
also with the pairs secured from laboratory (camp) mating. Those taken
July 16 and 17, 192 1, and August 16, 1922, always revealed this type of em-
brace. Our notes record that in one of the spadefoot pools a male spadefoot
had a ripe female Hyla gratiosa in the embrace. The male was the larger in
size. Unfortunately we have no memory of whether the amplexation was
inguinal or abnormal.

OVULATION

Habitat. We have found the eggs or tadpoles in open ponds of cut-over
pine lands, in cypress ponds open at one end and shaded on the other or in
dense cypress and gum ponds. At times in preference to nearby cypress ponds
we have found them in temporary grassy pools (after hard storms of 2-4 or
5 inches) such as spadefoots frequent or open tussocky transient pools beside

304 Frogs of the Okefinokee Swamp

the road. All in all it is primarily a mainland tree frog with preference for
cypress or gum ponds not so dense as the cypress bays or the ponds of
some islands. It incHnes to more open cypress ponds or cutoff cypress ponds.

Period. In 191 5 Deckert (p. 5) writes "I have never heard this species
call except during the breeding season which lasts from March to June, be-
ginning with the first warm rains." In 192 1 we were in the swamp from last of
April to September but in the center the species is scarce. In 1922 we were at
Chesser Island from mid- June to mid-September. Based on these two
summers' experience in 1923 (p. 34) we wrote "From June ist to loth three
species begin (to breed), one of which was Hyla gratiosa.'" In 1924 (p. 378)
we gave "Season June 10 to August 21." The few ovulations in summer in
192 1 and 1922, and its breeding after a severe storm in August 1922 with
Rana aesopus and ScapMopus holbrookn spring breeders led us to conclude it
to be a spring breeder with some tardy ovulation in summer to September i ,
The period seems March to August. One might suspect that the late breeders
were younger breeders. This is not always the case. Since this was written
we have noted the observations of Prof. T. Van Hyning (1923, p. 68) on these
very three species. He writes: "On the night of March 18, while some of the
boys of the biological class of the University of Florida, Gainesville, Fla.,
were collecting frogs, among other species taken, were thirty specimens of the
Florida Tree Frog, Hyla gratiosa LeConte; . . . These and other species were
all in a pond breeding near the University. One of the party collected eggs,
and brought living specimens which have spawned since, and is keeping records
on the incubation and development of each species.

"As there seems to be little really known concerning the breeding, es-
pecially of the . . . Florida Tree Frog, I consider the incident worth recording."

Temperature and humidity. (See voice for discussion of rain and tempera-
tures of choruses). On the dates when ovulation or mating occurred the
maxima averaged 90° and minima 71°. They were always preceded by heavy
rains.

Egg-laying process. We did not see the process in our captive pairs. In
nature Deckert (19 15, p. 5) says "The eggs are laid singly and sink to the
bottom of the pool." Whether they swim from place to place on the bottom
ovulating and fertilizing one egg at a time, or whether they ovulate and
fertilize several or one at the surface and let them or it sink needs to be ob-
served. The eggs are certainly disposed on the bottom.

EGGS

Attachment, egg )nass. On July 17, 192 1, about 11:00 a.m. we "went
to Hyla gratiosa pond. Cypress trees scattering with hurrah bushes (Leucothae)
around them. In amongst iris, fern {Woodwardia broad leaf), tussocks of
sedge found eggs. In water 3-6 inches deep with dip net discovered eggs
which seemed those of Pseudacris ocularis; later single eggs presumably
(Acris) with citrine upper poles and yellowish vegetative poles; and one or
two strings 4-6 eggs of Bufo quercicus. The supposed Acris eggs are loosely
attached to sedge or grass blades or oftener to sphagnum moss on the bottom.

Hyla gratiosa 305

They are easily seen. The Pseudacris eggs are harder to see." "None of these
eggs are H . femoralis eggs. I was careful not to scoop in surface packets which
were present. It is possible the yellowish single eggs are not of Acris but of
Hyla gratiosa or some other frog yet unknown."

On the following morning one of the mated pairs had laid a large comple-
ment. It filled the pan and was continuous. The pair had no escape and
could not strew them singly. Thereafter and the next year whenever there
were congresses of this species we never found a continuous film of their eggs
on the surface of the water as this check would lead us to expect. It was laid
in abnormal surroundings. We compared these positive eggs of our laboratory
pair with the supposed single eggs of Acris taken in pond the day before.
The supposed Acris eggs were Hyla gratiosa eggs.

Egg description. The measurements of 24-29 eggs give the following re-
sults: vitellus 2 of i.o; 8, 1.2 mm.; 12, 1.4 mm.; 6, 1.6 mm.; i, 1.8 mm.; range
1. 0-1.8 mm.; mode 1.4 mm., average 1.37 mm. The outer envelopes were i
of 2.3 mm.; i of 2.4 mm.; i, 3.0 mm.; 3, 3.8 mm.; 10, 4.0 mm.; 2, 4.6 mm.; i,
4.8 mm.; 5, 5.0 mm.; range 2.3-5.0; mode 4.0 mm.; average 4.06 mm. In the
field we measured the vitelli as 1.2-1.3 mm., and thought there was an inner
envelope. It is, however, the vitelline membrane far from the vitellus and
appearing as an inner envelope 1.6-2.0 mm. in diameter. The animal pole
is light brown or better olive in preserved eggs and the vegetative pole is
pale or yellowish. In nature it is above citrine or greenish brown and yellowish
below.

We have not counted the ovarian egg complements of captive females. On
July 23, 1922, we counted 1/4 of the area of the photo of the complement of
July 18, 1921, and secured 521. This gives a complement of 2084, a fair
number for the complement of an average female of this largest tree frog in
eastern United States. The female which laid this 2084 complement is 59
mm.; our largest female specimen is 68 mm.

Dangers. In general they choose cypress ponds. In that case the eggs
doubtless hatch before any danger from drying up. The males caught June
5, 192 1, July 3, 192 1 and pairs taken August 16, 1922 were from a corn field,
a pool by a wood pile and a shallow grassy temporary hole in the pine woods.
Every place would dry up in a few days at the most. The loss of tadpole life
would be great. No doubt this species suffers in this regard. It grows to the
largest Hyla size before transformation and the toll is surely great in this
species.

HATCHING PERIOD

The only record we have is from the pair caught the evening of July 17,
192 1. This pair laid eggs between 12:00 a.m.-6:oo a.m., July 18. On July 20,
we make the following note "No. 553 Hijla gratiosa eggs just hatched or better
night of July 19-20. Eggs laid in camp." This means two days from laying
in camp and in shallow water i inch or less. In nature they were in water 3
inches-i foot in depth.

3o6 Frogs of the Okefinokee Swatnp

MATURE TADPOLE

Color description from life {June 23, 27, A ugust 21 , 1922) . They have con-
siderable greenish, greenish yellow or yellowish in the tail and body and are
beautiful tadpoles. Half grown tadpoles have a striking black saddle spot
on the back of the muscular part of the tail and two prominent light areas
from each eye to the vent.

On the side a patch just ahead of developing hind legs light greenish yellow.
Black stripe from between the eye along the base of the upper crest on either
side half way to tip of the tail. Light area beneath this black area and around
nostril, over eye, along side to base of muscular part of the tail ivory yellow,
tilleul buff or pale pinkish buf¥. Another short hne of the same color at the
very base of the upper crest and above the black stripe. Muscular part of the
tail and crest yellowish citrine, light yellowish olive, mignonette green or
courge green. Belly pale vinaceous pink. Either side at gill region congo
pink, throat clear. Lower lip like muscular tail, yellowish citrine or courge
green.

General appearance. Tadpole medium (50 mm.) largest of Hylid tadpoles
of the eastern United States. Tail long. Tip acuminate, with flagellum.
Dorsal and ventral crests about equal either equal depth to the tail mus-
culature. Dorsal crest extending on body to a vertical about half way be-
tween the spiracle and the eye. Spiracle sinsitral, far below lateral axis,
directed upward and backward. Eye on lateral axis, in dorsal aspect in the
lateral outline and in consequence visible from the venter. Anus dextral, on a
level with the lower edge of the ventral crest. Muciferous crypts indistinct.

Mouth parts: Teeth 2/3. Upper labium fringed with row of teeth; the
papillae extend above and inwards beyond the end of this fringe 2/7-1/4
(usually at least 1/4) of the length of the fringe. The end of the second row
about even with the end of the fringe. Sometimes the ends unite. The horny
beak contained in the length of the upper fringe 1.5- 1.7 5 times. Median space
between the two lateral parts of the second rows contained 1.5-3.0 times either
lateral row. Inner papillae on either side from end of lateral upper row one or
two rows to end of lower third labial row. The lower third labial row with
one row of papillae below it, rarely with none, about equal to this single row
of papillae, or 2 5/8-3 1/2 times in the length of the first or second rows. The
first or second lower labial rows about equal or about i i /3 longer than horny
beaks. Some of the teeth rows have tendency to be curved on ends or united
at other rows or be quite irregular. More so than in any other United States
Hylid.

Measurements. Length of body (13.5-20 mm.) in tail (27-32 mm.) 2.3-
3.25, average 2.5. Width (9-13 mm.) of body in its own length 1.5-2.0, aver-
age 1.7. Depth (9.5-13.5 mm.) of body .82-1.0 in body width, average .9.
Depth of body 1.4-1.9 in body length, average 1.62. Depth (10-14 mm.) of
tail in length of tail 2.0-2.8, average 2.34. Muscular part (4.5-8.5 mm.)
1.5-2.9 in depth of tail, average 2.0. Spiracle i. 0-1.375 nearer base of hind
legs or vent region (8.0-10.0 mm.) than the tip of the snout (10-12 mm.),
average 1.2. Spiracle nearer eye than base of hind legs or vent, average 1.9.

Hyla gratiosa % 307

Eye 1. 07-1. 75 nearer to spiracle (4.0-5.6 mm.) than to tip of snout (6.0-7.0
mm.), average 1.3. Nostril i. 0-1.8 nearer eye (2.5-4.0 mm.) than snout (3.5-
4.5 mm.). Mouth (3.5-4.5 mm.) usually 1-1.5 larger than internasal space
(3.0-4.0 mm.), average 1.075. Mouth contained 1.33-2.4 (average 1.88) in
interorbital distance (6.5-8.5 mm.). Internasal space contained in interorbital
space 1.62-2.33, average 2.15.

The dimensions of the largest tadpole are :

mm.

mm.

Total length

50

Spiracle to vent

10. 0

Body length

19

Spiracle to eye

5-5

Body depth

12

Eye to snout

7.0

Body width

10 -5

Eye to nostril

4.0

Tail length

31

Nostril to snout

4.0

Tail depth

II .0

Mouth

4-5

Musculature

of tail

7.0

Interorbital distance

7.0

Spiracle to snout

II • 5

Internasal distance

3 5

General remarks. In 1921 we secured nothing on the larvae. In 1922 we
"found small tadpoles of this form in the pond i mile south of Trader's Hill.
They have a striking saddle back spot on the muscular part of the tail and two
prominent light areas from each eye to vent." On July 26 "In Anna's Pond
found large mature tadpoles of H. gratiosa. They are large creatures. They
have considerable greenish in the tail and body." On July 28 "in a pond
(open) I mile east of Chesser School mature tadpoles. The tadpoles were in
the drifting masses of vegetation." In 1923 (p. 406) we characterized the
tadpoles as follows: ''Hyla gratiosa — beautiful green tadpoles, deep crests,
young with black saddles on base of the tail, light line from eye to tail."

These long-tailed, big-eyed, at times flagellum-tipped tadpoles show the
myotomes of the tail musculature particularly along the edges of the mus-
culature. One captured July 31, 1922, has a forked tail-tip. Tadpoles about
half-grown, if viewed from dorsum, look to be a perfect wedge from eye to
tail tip.

LARVAL PERIOD

In 1922, in April at St. George and Way cross there was only .55 and .69
inches rainfall for the whole month, but in May 8.70 and 7.38 inches respec-
tively. But at St. George and Waycross 6.05 and 4.12 inches of the month
rain was from May 24-31. The transforming frogs of July 25-28 were pre-
sumably from this May period or 55-65 days as larvae. If, however, they
came from eggs laid in the great June 10-14 choruses which we found on
arrival, the period is reduced to 41-48 or 50 days.

We inchne toward May 24-31 as the period of ovulation because in this
pond the tadpoles which transformed July 26-31, 1922, were recorded as half
grown in this pond on June 27. June 10 or 12 is too early for these tadpoles,
better May 24 onwards or a month for growth to the half way period. This
species has to grow to a larger transformation size than any other Hyla in the

3o8 * Frogs of the Okefinokee Swamp

swamp and, therefore, would not be expected to have a short period, but
rather a long one. The upper range 41-65 may be a minimum period of
development.

TRANSFORMATION

Period. In 1922 on July 10 ''found 2 or 3 tadpoles in two-legged condi-
tion." On July 17 they were not beyond the two-legged stage. On July 24
"found one two -legged tadpole ready to burst out its forelegs," and on July
25 "the tadpole burst its forelegs this morning." On July 27 we found this
tadpole fully transformed. Two days later at same pond found four trans-
formed or transforming. On July 26 in another pond found a transformed
frog. A month later, August 21, 1922, in a third pond "Several beautiful
mature Hyla gratiosa tadpoles. One a beautiful specimen with a very
yellowish or greenish yellow (non-Ridgway) colored tail. In one pond we
record on August 9 that the Hyla gratiosa tadpoles are quite small. All in
all we actually took transformed individuals from last of July to the last of
August.

On July 26, 1922, when coat bets were having a great breeding congress
where we solved the "barker" puzzle we found breeding signs and "found a
transformed Hyla gratiosa 2 or 3 inches above the water on a spear point leaf
of Equisetum (horsetail)."

On July 31, 1922, we have these journal observations. "In one pond we
found in one haul of the seine 6 Notophthalmus dorsalis larvae, Hyla gratiosa
mature tadpoles, transformed frogs and a tadpole with bifurcated tail.
Plenty of Hyla gratiosa and Rana aesopus tadpoles. Caught two Kinosternon
subrubrum. In a pond south of this first pond took almost a pure culture of
Hyla gratiosa tadpoles with a few red-tailed Hyla femoralis tadpoles. In
pond (corner of Chesser Island and Old Okefinokee roads) secured largest R.
aesopus tadpole and plenty of Hijla gratiosa transformed and tadpoles
a plenty." Thus the progeny of our earliest breeders may be transformed or
transforming when the later breeders are just beginning. Eggs laid in July
or as late as August 16 must transform in September or October. We thus
have July-October at least for transformation.

Size. On July 26, 1922, a transformed frog measured 20 mm., on July
27 three were 19, 23, 21 mm, respectively; another of the same date measured
19 mm.; on July 29, two measured 18 mm. and two 20 mm.; on July 31 we
have three 19, 20, 21 mm. in length and on August 21, one of 23 mm. We
have only 13 specimens with an average and mode of 20 mm., and a range of
18-23 mm. This makes it a larger transforming Hyla than Hyla versicolor
with its 14-20 mm. range.

General remarks. " Hijla gratiosa when transformed often shows brownish
spots on the back; sometimes has three or four yellow or orange spots like an
adult Hyla ci?ierea. Dark vitta from eye backward and three or four dis-
connected dark spots on sides almost to groin. Another shows the spots of
dorsum common and dark green."

As transformation approaches the myotomes of the tail musculature show
very clearly. The creature looked like a big-eyed, big-headed creature with

Hyla gratiosa 309

very slender waist and hind legs, the head being very broad. The body looks
very cuneate from eyes to waist to tip of tail. At transformation the tym-
panum may not be evident at all. The granulations of the dorsum begin to
show before transformation is completed.

GROWTH

At Hilliard, Fla., August 16, 1922, we took males 55, 59, 60 mm. respec-
tively, two groups (?). At Trader's Hill on July 4, 1922, we secured two males
52, 56 mm. On Billy's Island in 192 1 we secured males 60, 60, 61. From
July 15-17, 192 1, Chesser Island, we secured 58, 61, 63, 65, 65, 66, 68 mm. cf s,
59, 68 mm. 9 s. Our material apparently shows two groups 52-56 mm. and
58-68 mm. with transformation size at 18-23 nini-

A glance over the U. S. National Museum material reveals that Dr. T. H.
Bean (U. S. N. M. No. 4742) at Micanopy, Florida, secured two groups 28,
30, 31, 31, 32, 32, 33-5 mm. and 47, 47, 49 mm., and Major J. LeConte (5901)
took 51 mm. cf, 56 mm. cf, 56.5 mm. 9 and 60.5 mm. cf and 9. There
seems three groups 28-33.5 mm.; and 47-56 mm.; 60.5- . In 'some

ways the evidence seems 18-23 mm. at transformation, 28-34 mm. first year
olds; 37-45 mm. second year olds; 47-56 mm. third year olds; 56-68 mm. four
year olds.

FOOD

There is little of record regarding the food habits of this species.

ENEMIES

On July 24, 1927, at Anna's Pond 15-20 feet in diameter found "much
fewer tadpoles than before. ... A Green Heron flew up. Twice a chicken
turtle (Deirochelys reticularia) has been seined out of it and a black snake,
watersnake and riband snake taken around or in it. All doubtless take their
toll of tadpole or transformed frogs." On July 27, 1927, we found 5 or 6
transformed Hyla gratiosa, one with the hind foot gone." A female of a pair
captured July 17, 1921, on Chesser Island, had its left forearm and foot gone,
and in the front of the right thigh is a sharp cut gauge as if bitten by a turtle
or some animal.

AUTUMNAL DISAPPEARANCE

Deckert records it in March- June at Jacksonville, Fla. We at Okefinokee
Swamp a short distance away have recorded it from June to September i.
The U. S. National Museum have records of specimens taken Sept. 9, 1912,
and Nov. 20, 19 12. The known range is March to November 20. It may be
more or less active the year through, though seldom seen or heard except at
breeding time.

AFFINITIES

This Sabalian frog, like some others has outgrown the ''Florida" handle.
The singly deposited eggs may place it with Hyla squirella, Acris gryllus,
Pseudacris ocularis and Hyla crucifer. This I would not emphasize so much

3IO Frogs of the Okefinokee Swamp

as the strewing of eggs on vegetation on bottom. This suggests more especially
H. andersonii, Hyla squirella, Hyla cinerea.

On June 26, 1922, when we found numerous mature tadpoles we entered
the following note "In relationships, on appearance H. gratiosa, H. cinerea
and possibly H. squirella may be closer much as Hyla femoralis seems very
close to Hyla versicolor." In a synopsis of tadpoles on mouth part characters
we have associated the tadpole of Hyla gratiosa with those of Hyla andersonii
and Hyla cinerea. In some ways the adult belongs nearer these two than any
others. Yet the size of the tadpole suggests the larger tadpole class size of
Hyla versicolor, Hyla arenicolor and Hyla femoralis. A perfect adult tadpole
has a flagellum-like tip like some of these forms.

BIBLIOGRAPHY

1889 Cope, E. D. The Batrachia of North America, pp. 379.

1915 Deckert, R. F. Copeia, May 15, 1915, No. 18, p. 5.

1906 Dickerson, M. C. The Frog Book, pp. 124-126.

1856 Le Conte, John. Froc. Phila. Acad. Nat. Sci., Aug. 1856, p. 146.

1922 Loding, H. P. Geol. Surv. Ala. Mus. Paper N0.5, Sept. 1922, p. 19.

1923 - Stejneger, L. and T. Barbour. A Check List of North American Amphibians and

Reptiles. 2nd edition Cambridge, 1923, p. 31.
1923 Van Hyning, T. Copeia, May 20, 1923, No. 118, p. 68.
1923 Viosca, Percy, Jr. Copeia, Feb. i, 1923, No. 115, p. 37.
1923 Wright, A. H. The Anatomical Record, January 20, 1923, Vol. 24, Number 6, p. 9, 406.

1923 . Copeia, Feb. i, 1923, No. 115, p. 34.

1924 and A. A. Wright, American Naturalist, July-August 1924, Vol. LVIII, No.

657, P- 378.

Hyla squirella Latreille

(PI. II, Fig. 7; V, Fig. 4; VII. Fig. 5; X, Fig. 7; XII, Fig. 9; XV, Fig. 5; XVII; XXXI;
XXXII; Text Figs, i, 10)

COMMON NAMES

Southern Tree Frog. Squirrel Tree Frog. The Squirrel Tree-Toad. South-
ern Tree Toad. "Scraper Frog." "Rain Frog." Squirrel Hyla. Tree Frog.

RANGE

Check list. "Type locahty: Carohna. Range: Virginia to Florida, west
to Texas, and northward up the Mississippi basin to Indiana." — Stejneger
& Barbour Check List (1923, p. 31).

Supplementary records. Deckert in May 1922 (1922, p. 88) in Dade Co.,
Florida, "saw several in Hammock on shell mound about 5 miles northeast
of Cape Sable. Heard their calls at Royal Palm Hammock." Myers (1924,
p. 60) reports "several" Hyla squirella received from Wilmington, N. C.
Schmidt (1924, p. 68) has it from Mt. Pleasant near Charleston, S. C. Holt
(1924, p. 9s) writes " Hijla squirella Latreille. Recorded by Loding from
Mobile County only. I have taken the species at Barachias, Montgomery
County."

Local Okefinokee records. In 1912, 1913 and 19 14 we did not take this
form in the swamp. In 192 1 we did not secure it until we went to the eastern

Plate XXXI
Squirrel tree-frog {Hyla squirella)

1 . Male (brownish individual) croaking, Anna's pond, Folkston, Ga. July 3, 1922.

Flashlight.

2. Male croaking. Camp Pinckney, Ga. August 11, 1922. Flashlight.

3-9. Tadpole series from mature tadpole to transformation, Anna's pond, Folkston,
Ga. June 27, 1922. x i.o.

10. Transformed frog. Ventral aspect, x i.o.

11. Transformed frog, Chesser Id., Ga. June 29, 1922. Lateral aspect, x 1.0.

12. Half-grown frog, Chesser Id., Ga. June 29, 1922. Lateral aspect, x 1.0.

13. Male, Camp Pinckney, Ga. June 21, 1922. Ventral aspect, x 1.0.

Hyla squirella 311

edge of the swamp in the last of July. In August of the same period it was
heard several times along the Suwannee and St. Mary's Rivers.

In 1922 from June 10 to September this species was recorded. Our opera-
tions were mainly outside the swamp. We have not taken it on any of the
islands of the swamp except Chesser Island.

GENERAL APPEARANCE

Brimley (1907, p. 154) gives this frog with "Size small, feet not edged with
yellow. Yellow or white band on sides not sharply defined above and below.
Back of thigh not marked with yellow spots or variegations. Dickerson
(1906, pp. 149) calls it a delicate, smooth-skinned little frog." Some have
alluded to the constancy of the transverse bar between eye or a part of it.
This sometimes disappears. Even the white on upper lip may be rarely
absent. Then the general appearance of the creature is about all one has in
identifying it.

MEASUREMENTS

(Recent Material)

i/eof/ to angle of mouth 2.85 (20 mm.)-3.i (28 mm. cf)-3.3 (28 mm. 9)
-3.27 (36 mm. 9) in length of body; head to rear of tympanum 2.85 (20 mm.)
-2.7-2.8-2.88 in length of body; head to angle of mouth 1.0-1.1-1.23-1.13
in width of head; head to tympanum 1.0-.95-1. 05-1.0 in width of head;
snout .71-.8-.7-.70 in first finger; snout .71-1.0-.9-.92 in fourth finger; snout
.42-.4-.8-.61 in first toe; eye .6-.5-,66-.75 in tympanum; eye 1.0-1.0-1.15-
1.12 in first finger; eye i. 4-1. 25-1. 66-1. 62 in snout; tympanum 4.0-4.25-4.0-
3.3 in intertympanic width; tympanum 2.3-2.5-2.5-2.16 in snout; internasal
width 1.0-1.5-1.0-.85 in upper eyehd width; interorbital width .5-1.0-1.0-.75
in upper eyelid width; interorbital width .5-.66-1.0-.87 in internasal width;
interorbital 2.0-2.8-2.66-2.5- in intertympanic width.

Forelimh. Forelimb 1.66-2. 15-2.07-2.05 in length of body; forelimb
2.75-3.0-2.8-3.1 in hindlimb; first finger i. 6-1. 375-1. 85-1. 55 in third finger;
second finger i. 6-1. 375-1. 55-1. 16 in third finger; second finger 1.0-1.0-.75-
.87 in first finger; third finger .7 5-. 7 7-. 8 5 in second toe; fourth finger 1.6-
1.1-1.44-1.16 in third finger; fourth finger .6-.4-.88-.66 in first toe; inter-
nasal width 1.6-2.0-1.16-1.28 in first finger; internasal width i. 6-2. 0-1.33-
1.7 in second finger; internasal width 2.6-2.75-2.16-2.00 in third finger,
internasal width 1.6-2. 5-1. 5-1. 7 in fourth finger.

Hindlimh. length 1.6 5- 1.4- 1.3 5- 1.5 in hindlimb; tibia i.8-i.8-( )-
( ) in length; tibia 3.o-2.5-( )-( ) in hindlimb; tibia i.i-.8-( )-
( ) in forelimb; tibia .63-.7o-( )-( ) in hind foot; first toe 2.0-1.75-
1. 25-1. 5 in second toe; first toe 3.33-3.5-1.87-2.2 in third toe; first toe 4.0-4.5-
2-37-3-oinfourth toe; first toe 3. 3-3. 5-1. 75-2. 12 in fifth toe; second toe 1.66-
2.0-1.5-2.25 in third toe; second toe 2.0-2.5-1.9-2.0 in fourth toe; second toe
1. 66-2.0-1. 4-1. 41 in fifth toe; third toe i. 2-1. 28-1. 26-1. 33 in fourth toe;
third toe 1.0-1.0-.93-.94 in fifth toe; fourth toe 1.16-1.2-1.9-1.75 in hind foot;
fourth toe i.8-i.7-( )-( ) in tibia; fourth toe 2.0-1. 44-1. 42-1. 49 in

312 Frogs of the Okefinokee Swamp

forelimb; fifth toe i. 2-1. 28-1. 35-1. 4 in fourth toe; internasal width i.o-i.o-
1.15-1.14 in first toe; internasal width 2.0-1. 75-1. 66-1. 7 in second toe; inter-
nasal width 3.3-3.5-2.5-2.57 in third toe; internasal width 4.0-4.5-3.16-3.4 in
fourth toe; internasal width 3.33-3.5-2.33-2.42 in fifth toe.

HABITAT

LeConte (1825, p. 279) says it "Inhabits under logs, and the bark of decay-
ing trees; I have never seen it in, or near the water." Holbrook (1842, Vol.
IV, p. 124) holds "This animal is found on trees, often seeking shelter under
the bark of such as are decaying; it frequently chooses old logs for its place of
hibernation. Often I have found it about old houses, and under logs and
boards." Deckert (19 14, No. 3, p. 3) at Jacksonville 191 1 and 19 12 found
them "in gardens and corn fields." The same author (19 15, p. 3) in his
Further Notes on Salientia of Jacksonville, Fla., writes: " Hyla squirella
Bosc, the commonest of the southern tree toads, is found everywhere, in
corn fields, sugar cane, about wells and under eaves of stable roofs, barns, out-
houses, etc. After a heavy thunder shower in September, 191 1, hundreds of
these little tree toads could be seen hopping along the country roads outside
of Jacksonville. That same night the writer took 32 specimens from a velvet
bean vine on the backside of his cottage." We found these frogs on porches,
in China berry-trees, oaks and other trees, as well as vines around the houses,
in fields and gardens around buildings, in open ponds, in pine barrens, in
pine and oak groves, along roads, in shallow roadside and pine barren pools.
On June 22, 1922, a boy at Camp Pinckney brought me a Scraper taken in his
own house.

Viosca (1923, pp. 37, 38, 40) found this species in the alluvial ridges of the
lowlands of southeastern Louisiana. "The fauna of these ridges differs least
from that of the uplands and there are many species in common. Some forms,
such as Hyla squirella . . . are typical .... Where the beach connects with some
alluvial ridge, oaks, bushes, and other ridgeland forms make up the vegetation
and the fauna partakes of the nature of alluvial species Hyla squirella and
Gastrophyne carolinensis sometimes reaching to the very edge of the Gulf."

FIRST APPEARANCE

Our earhest record is June 10. In Dade County, Deckert records them in
May and June. At Houston, Texas, Pope (19 19, p. 90) reports "This species
was more abundant than any other Hylas, but was the latest to appear in the
spring. The first one was collected April 13." The U. S. National Museum
has specimens from L. Tohopikaliga, Osceola Co., Feb. 6, 1906, Biloxi, Miss.,
Feb. 16, 1914, Dauphin, Ala., Mar. 9, 1892, and from Lake Kissimee, Osceola
Co., Fla., Apr. 3, 1901. We have specimens from Bay St. Louis collected in
January.

GENERAL HABITS

Metachrosis. This very apparent phase of tree frogs has been discussed
more than any other topic. This species is a capital example of the sudden
changes of which Hylas are capable.

Hyla squirella 313

LeConte (1825, p. 39) writes of it at length: "The figure of this species in
Daudin is very different from any specimen that I have ever seen, both in
colour and in marks ; I do not deny but that it may have been fond of a green
colour, as he represents it, and marked in the same manner; but the same error
occurring in his delineation of the next species, I am incHned to think that it
has been coloured from report and description, rather than from the animal
itself; indeed, whatever the colour may be, it can scarcely remain unchanged
for six months, in a preserved specimen; all that I have ever attempted to
preserve, lost their colour in less than two months. There are three principal
varieties of this species:

"a. Above cinereous, with a straight, or curved or angular bar between
the eyes; hack with a few spots of dusky, sometimes confluent, and form-
ing different figures of irregular shapes; and sometimes uniting into a
line on each side of the body, of greater or lesser length.

'*b. Above cinereous, irregularly spotted with darker; the line between
the eyes broken into two or more spots.

"y. Above entirely brown, with spots, exterior part of thighs not
yellow.

"In all these, the dark band on the head, and the white line on the hps,
are the only marks which remain constant; in one variety, even the yellow
colour on the thighs vanishes; there are hardly any two individuals alike, and
so different are they from one another, that a person who had not observed
them accurately for a length of time, would be led to think, that there were
almost as many species as individuals."

Holbrook (1842, p. 124) remarks that "The colours of this animal are even
more changeable than in any species with which I am acquainted. I have seen
it pass in a few moments from a light green, unspotted and as intense almost
as that of Hyla lateralis, to ash-colour, and to a dull brown with darker spots;
the spots also at times taking on different tints from the general surface.
The markings, too, vary exceedingly in different individuals, the white line
on the upper hp and the band between the orbits alone are constant."

LeConte (1855, p. 429) again writes of it as follows: "Color varying at the
will of the animal from green to brown of different degrees of intensity,
spotted and speckled irregularly with darker and dusky and sometimes with
paler, often however of a uniform color. A darker line extends from the
nostrils to the eyes and through them to the insertion of the arm (this is some-
times evanescent) ; beneath this darker fine extends a white one which reaches
nearly to the groin; sometimes interrupted or broken into three or four parts.
The dark line on the vertex between the eyes never fails entirely, although it
is sometimes reduced to a rather large spot on each eyelid."

Dickerson (1906, pp. 149, 150) remarks this wonderful change of colour
the "scrapper" shows. "Of all the tree frogs of North America, this one has
perhaps the greatest power for rapid colour changes, and during these changes
presents the greatest variety of colours and shades of colour. At any given
moment, Hyla squirella may wear any one of the following costumes: Un-
spotted dark chocolate brown or dark brownish olive; light purplish brown

314 Frogs of the Okefinokee Swamp

with dark brown spots; light yellowish or greyish brown without spots; any
medium shade of brown with green spots; olive green unspotted; light yellow
green spotted with brown; unspotted light pea-green; light greenish gray;
light fawn colour, or still lighter shades ranging down to flesh colour."

"Most curious is the fact that although these changes take place under the
influence of various stimuH, such as light, moisture and heat, they may go on
without these stimuli. Frogs shut in a dark pail with no change of conditions
will not appear twice alike when the pail is opened at intervals during the
space of several hours. Some may be green and others brown; some spotted,
others not; some light while others are dark. And at any given time of
observation those that were dull and spotted before may be a light, unspotted
green, the ones that were Ught may be dark, and so on. The light line along
the jaw undergoes great changes also. This tree frog furnishes a most interest-
ing case for the study of metachrosis."

VOICE

Deckert (191 5, p. 3) at Jacksonville, Fla., describes its cry as follows:
"The cry is rather coarse, sounding like 'era, era, era', etc., with a second's
interval between each note." The same author (192 1, pp. 22) in Dade County,
Florida, "Their rasping calls were heard in May and June at 19th Street,
Miami, at Donn's nursery and at 22nd Street in company with the toads
mentioned previously. After heavy rains in July, August and September
they were heard also at Cocoanut Grove, Lemon City, Little River, Arch
Creek and Fulford."

Pope (19 1 9, pp. 96, 97) at Houston, Texas, gives considerable details of
its calls. "The call is a harsh, rasping trill not so loud nor so musical as that
H. versicolor. From a distance the chorus resembles that of H. crucifer, but
the call is much harsher when heard close at hand. The throat pouch of the
male is large and expands to nearly the size of the body. . . . July 31. Have
heard a few frogs almost every night lately, calling from trees near sleeping
quarters. They do not begin to call until ten or eleven o'clock at night, a^nd
may continue intermittently till daylight. Usually a single one begins to call
and is joined by half a dozen or so more until there is quite a chorus."

The first squirrel tree frog the writer ever heard came June 10, 191 7
beyond Tuskagee, Ala., and 30 miles from Montgomery. We went through
a fine series of lowland woods and over several wooden bridges. "Here first
saw Sabal serenoa. Heard Whiteeyed Vireo, Acadian Flycatcher, Carolina
Wren, Hooded Warbler, Southern Parula Warbler, Tufted Titmouse, Caro-
lina Chickadee, etc. ... In a moist field heard no end of Acris gryllus. After
dark heard another note which proved a male Hyla squirella. Took one on
top of a bush 8 feet high. Another we supposed high in a tree although we
found it on a bamboo-like cane 5 feet from the ground."

In 192 1 we first heard them on July 21. "Probably the tree frog we heard
in Chinaberry tree at Mr. Rider's house during the storm was H. squirella,

Hyla squirella 315

the Scrapper Frog. Mr. Rider avowed that they were all around and that
they can change color to suit their environment. Some of them he maintains
have no conspicuous marks."

Harper made the following notes on this species on his trip in August on
the Suwannee and St. Mary's Rivers: "At Fargo a number were heard during
a hard rain on the afternoon of August 5, and one was found on a potted
prickly pear (Opuntia) on the porch of a house. Here it was occasionally
giving its rather weak and leisurely diurnal note: Waaaake, waaaake, etc. It
has a sort of grating or scraping quality and might well be compared to the
scraping of a hollow reed with a fine-toothed comb. It is probably not to be
heard at a distance of more than 50 or 75 yards. In the evening of the same
day a big chorus was heard at Moniac.

"On August 14 and 15, in late afternoon and early morning, several were
heard in a pine grove on the St. Mary's River (Georgia side) about five miles
south of Traders's Hill. On August 15 and 16 several were heard in damp
woods on the south side of this river near Boulogne, Fla.

"During a thunder shower on the afternoon of August 16 a number began
calling at Camp Pinckney, and one was captured among the leaves of a live
oak about 15 feet from the ground. After dark numbers were on the ground,
especially in or about a temporary pool, in a road through level oak woods,
where their chorus was deafening. The nocturnal note is similar to the di-
urnal one, but stronger, louder, and more regular; in fact, it is almost a con-
tinuous call. I found it being given at the rate of 90-95 times per minute.
In the alternate inflation of throat and body, the throat sac attains about
half the size of the body. When sitting on the ground, croaking, this frog has
more or less the appearance of a large, fat Acris. Some individuals were
green, and others, I think, brown.

"On the morning of August 18 several others were heard farther down the
river at Kings Ferry and vicinity."

In 1922 our first introduction to this species came seven miles north of
Millen, Ga. "Here at 7 p. m. we heard several and took two in the small
trees at the edge of a swampy tangle. Also heard them in higher trees. An
imminent storm made them noisy. The following morning we heard a few."

On June 19 "At Camp Pinckney hear a Hyla squirella across the St.
Mary's River on the Florida side. On the ground saw one hopping along.
We mistook it for a frog {Rana). It was brownish with spots. Leaped on to
some dead branches. It can leap. When the thunderstorm came several
Hyla squirella were to be heard and some H. versicolor. We captured one in
a tree near the sawmill." On June 23, near the old railroad pools near
Trader's Hill heard one or two scrapers in the afternoon. On June 26 at
Anna's Pond heard one calling. On June 2 7 at Camp Pinckney heard several
in mid-afternoon. "On a fence heard a Hyla squirella. A little later heard
another frog note from a fence where the H. squirella was. Were they the
same? Is it versatile in its calling? Later a thunderstorm threatened; heard
several Hyla squirella."

3i6 Frogs of the Okefinokee Swamp

On July 2 at ii :oo p. m. heard only one or two near the house. On July
3 we heard them along the Folkston Road. "At 7 115 p. m. we started for the
hill. We began to hear H. squirella on our way back from the Trader's Hill
ponds (i mile south) and came to a pond 1/2 mile west of Trader's Hill. It
was a grassy overflow pool. In a clump of bushes and saw palmetto were
several Hyla squirella. In grassy shoals in shallow water were several, one of
which we photographed. In same grassy overflow were three or four Hyla
gratiosa calling, Hyla femoralis common, Bufo quercicus abundant, Acris
gryllus more or less present. This was at 9:50 p. m. There were Bufo terres-
tris present. Along on Folkston Road in road in temporary pools and ditches
with Bufo terrestris were plenty of Hyla squirella. At 9:30 I heard a Hyla
squirella give 67 pumps in 45 seconds. There is more vibration in the call of
Hyla femoralis. Could not hear the Hyla squirella a few rods away. The
calls of Hyla femoralis and Bufo quercicus drown it out. Hyla squirella does
sometimes croak from water surface when sprawled on the water."

On July 9 we heard only one at Thompson's Landing. On July 20, 3 miles
from Jacksonville, Florida, we heard another about 4:00 p. m. On July 24
heard a few in mid afternoon at Camp Pinckney. On August i several were
heard in late afternoon and evening at Waycross and Hebardsville, Ga. On
August 8 in mid forenoon we heard many near StarHng Branch. The follow-
ing day in Spanish Creek woods heard them during rain at noon. "As we ap-
proached Trader's Hill heard many Hyla squirella at 1:00 p. m."

On August II "at Camp Pinckney 2-3:00 p. m. heard several. At 8:30
returned to Camp Pinckney. No end of Hyla squirella. They were on the
ground in a road filled with temporary pools, water 1-3 inches deep. Count-
less males. Sac hyaline, more or less inflated for some time. Not so fast in
caUing as Hyla femoralis but swift nevertheless. Those in water greenish.
Those on edges of pools or in road brownish. . . . Could find no females.
Some males, though quite small, are croaking."

On August 13, on a trip from Chesser Island to St. Mary's, Ga., we heard
them in several places but especially from 7-10:30 along the Folkston Road."
From August 15-17 we heard several at Callahan and Hilliard, Florida, and
I or 2 at Moniac, August 20. Our last record of calling came September 15, 1922.

Temperature and humidity. In 192 1 the great chorus of "scrapers" came
August 5. They were calling during the rain or because of rainy weather.
The combined rainfall of three places around the swamp was 7.02 inches,
maxima 90-95°, minima 70-73°. Thereafter until August 18 a few were
heard, maxima 87-97°, minima 68-76°. On August 16, F. Harper and
Marion Lee found them calling vigorously from their tree homes in daytime
at Camp Pinckney when the thunder storm came on. After dark, numbers
were on the ground, especially in or about a wet place in the road thru oak
woods, where their calling was deafening. Act like Hyla cinerea, throat
large inflated, about half the size of the body. Scraper frog called 'Rain
Frog' by natives here about. One I timed gave about 15 calls in 10 seconds,
three times when I timed it and about 16 times in 10 seconds the fourth time.
A very regular and mostly continuous call.

Hyla squirella 317

In 1922 on June 19 at Camp Pinckney a severe storm came from i :oo to
3:00 p. m. Hyla squirella were calling vigorously.

On June 23, two inches of rain fell at St. George and scrapers were calling.
On June 26 a half inch to an inch of rain fell, and the Hyla squirella were
calling strongly at Anna's Pond on that day and on June 29 at Camp Pinckney.
On the afternoon of July 2 and through some of July 3 considerable rain fell
and the species were common in roadside ditches west of Trader's Hill, in
Anna's Pond, Starling Branch, Cornhouse Creek and several other places.
Another great chorus came August 16 and 17 after a half day's rain when
about two inches of rain fell, maxima 82-91°, minima 65-74°. Spadefoots
and Rana aesopus, subterranean species came out to breed and Hyla squirella
as well.

In general this species calls even by day in rain or before an imminent
rain (July 9). After a downpour of warm rain 1-5 inches they became very
active. As every observer has remarked heavy warm rains bring them out in
midsummer. During our 1922 study June lo-August 20, maxima were 81-
90°, minima 62-74°, averages 90°, maxima minima 70°.

MATING

Alale {Fi'om life, June 22, 1922). Back citrine or buffy citrine in one; on
another ecru-olive or light yellowish olive; rear back and top of hind legs
chrysolite green to lime green. Under parts of hand also hind foot orange
rugous, xanthine orange to orange chrome; hind Hmb posterior and anterior
faces and throat raw sienna or mars yellow; throat of another aniline yellow
or other hind leg parts raw sienna. Stripe on upper jaw on to body greenish
or lemon yellow. Belly cream color or ivory yellow. Tympanum hazel or
russet. Iris spotted black and army brown, a fawn in one deep chrome and
antique brown in another. A light area of greenish on either side of throat
but not clearly defined or definite as in Hyla andersonii and Hyla cinerea males.

One male held in hand in good light became courge green and another oil
green. A small last year's H. squirella was light dull-green-yellow above and
upper jaw stripe light chalcedony yellow. Sometimes those in water may be
greenish while those on edges or on land may be brownish.

Female {From life, July 3, 1922). Four females of four different pairs had
under parts white with no discolored throats. Throat slightly primrose
yellow, reed yellow or white. Forelimbs, groin, hindlimbs before and aft,
tibia and hind feet reed yellow, olive yellow. In one yellow ocher on fore and
rear part of femur, under side of tibia and foot and fore foot. The females
unless very large have none of the bright orange rufous, xanthine orange or
orange chrome of the males.

Structural differences. In the four mated pairs the pairs were in size as
follows: cT 31 mm., 9 30 mm; cf 29.5 mm., 9 28.5 mm.; cf 27 mm., 9 27
mm.; cf 28.5 mm., 9 35 mm. In other words in two cases the males were
larger, in one case male and female equal and one where female was much
larger. We have females 27 and 35 mm., and twelve males from one place
ranged from 27.5-35 mm., average 32.4 mm., mode 32 mm.

3i8 Frogs of the Okefinokee Swamp

These are about the maximum size. Of 92 specimens measured in the
U. S. National Museum there were only three of 30 mm., one of 31 mm., two
of 32 mm., one 33 mm., and one 35 mm.

Of a group of 35 specimens collected at South Jacksonville, Florida by
Thomas HaUinan in 1922 for the American Museum of Natural History there
are of females (26) two 23 mm., two 24 mm., five 25 mm., one 26 mm., three
27 mm., three 28 mm., one 29 mm., one 30 mm., three 31 mm., three 32 mm.,
one 33 mm., and one 36 mm.; of males (7), one 23 mm., two 24 mm., one 25
mm., one 28 mm., one 30 mm., and one 34 mm. ; and of doubtful but apparently
females one 19 mm., one 20 mm., three 21 mm., four 22mm., one 23 mm.,
and one 25 mm. One lot of three specimens (Am. Mus. Nat. Hist. Nos.
14968, 14969, and 14970) collected by Jay A. Weber, 192 1, at Ocalawacochee
Slough, Lee Co., Florida, are respectively 20, 23, 24 mm., but we record that
"they look like females but are they or are they immature?" This American
Museum material shows males from 23-34 mm., females from 23-35 mm.,
but, however, doubtful females from 19-25 mm. Sizes from 21-24 mm. do
not always reveal the sex externally. The extreme size of females found is
37 mm., and of males 36 mm. Han Geyer (1902 pp. 98, 99) reports that he
had one 45 mm. in length.

The female has the hght throat and the males the more intense throat.
In twelve males taken at same time in spirits none had a dark pigment on
throat except one which was heavily spotted with dark.

Duration. On July 3, 1922 "About 11:55 or midnight we were at Anna's
Pond. Heard the Hyla squirella at a distance between two distant houses.
In the saw palmettos and grass stools around the pond were many scrapers.
One pair found in grass near the edge. Other pairs: three were found in saw
palmettos near edge of pool as were the Hyla femoralis in pairs. The pair of
Hyla squirella we photographed hopped upon the road some 8 or 10 or more
feet from the pond. By 7 :oo a. m. the next morning all four males were not
mating. All four females had laid during the evening."

Night or day. Most large choruses come at night though severe rain
storms might bring breeding in day time. The frogs prefer the security of
night to come down to the ground.

Amplexation. — (Normal, abnormal). All the males of our mated pairs had
the axillary embrace. On August 11, 1922, at Camp Pinckney we make the
note "Once in a while a male chases another away or at times tries to mate
with it."

OVULATION

Habitat. We found them breeding in a roadside ditch, in an open saw-
palmetto-surrounded pond (once pine woods), in grassy pools in open land
and near a road. The tadpoles we found in some of the above places, and in
pools beside an old railroad embankment.

Period. Pope (1919, p. 96) at Houston, Texas, on "April 24, 1918, found
them breeding abundantly in a shallow pool after a warm rain." "May 4. A
heavy warm rain this evening brought out Hyla squirella in force. This
seems to have been the regular breeding season and for some time after I

Plate XXXII

Squirrel tree-frog (Hyla squireUa)
Anna's pond, Folkston, Ga. June 27, 1922.
Mated pair, Anna's pond, Folkston, Ga. July 3, 1922.
Egg mass, Hilliard, Fla. August 17, 1922.

5. 6. Male croaking, Trader's Hill, Ga. July 3, 1922. Flashlights
Female and male, Chesser Id., Ga. July 4, 1922.
Eggs, not stained, Chesser Id., Ga. July 4, 1922.

Hyla squirella 319

heard very few of these frogs except for a httle after every rain." We have
breeding in June, July and August but transforming frogs recorded from June
26- July 10 implies early spring or spring breeding is common. In 1923 (p. 34)
we held that "From June ist to loth, three species begin (laying). Bufo
quercicus. . . . Hyla squirella chooses the bottom for its singly laid eggs as
does Hyla gratiosa." In 1924 (p. 378) we give the season June lo-August 21.
Doubtless the bulk of laying in the Okefinokee country comes from April to
June because of the transforming tadpoles in June and July.

Temperature and humidity. Actual ovulation or mating in 1922 came once
after a 2 inch rain, maxima 87-96°, average 92°, minima 68-72°, average 70°;
after 1.42 rain; after 1.60 inch rain, maxima 81-93°, average 89°, minima
65-83°, average 72°; after 1.80 inch rain (August 16) maxima 82-91°, average
88°, minima 65-74°, average 71°. In combination we have rains i. 42-1. 80
inches, maxima 81-96°, averages 78-92°, minima 65-83°, averages 70-72°.

Egg-laying process. We did not see the egg laying of any of the four pairs
we had in captivity. Apparently they lay single eggs. Whether they are of
the Hyla crucifer type or Hyla andersonii type or like Hyla gratiosa we cannot
say. Whether they remain at the top and strew eggs singly {Hyla andej-sonii)
or several at a time, or whether they progress on the bottom like Hyla crucifer
and lay single eggs remains to be observed.

EGGS

Egg mass. Pope (19 19, p. 96) is one of the only workers to record their
eggs. On April 24, 19 18, at Houston, Texas, he "Caught one mated pair and
they laid eggs in a jar the next day. The eggs are laid singly, but tend to
cling loosely to the bottom of the jar and to each other."

We have in every instance thus far held that they were singly laid. But
on June 22 at Camp Pinckney we found in one of the water pits a surface film.
At first we naturally concluded they were Hylafemoralis eggs. Closer exami-
nation made of the individual eggs made us question "Are they H. squirella
or Hyla versicolor eggs? Later we found Hyla versicolor tadpoles in the pond."
Still they may be Hyla squirella. Our observations on ovulation in this
species are too few.

Egg description. Pope (19 19, p. 96) describes the eggs laid in captivity
for him as follows: "In color they are light brown above and nearly white
underneath. Diameter of egg not quite one mm., vitelline sac about .12 mm.
in thickness, jelly outside that .25 mm. thick."

On the night of July 3-4, 1922, "at Anna's Pond 11:55-12:00 or later we
heard the Hyla squirellas. . . . Four pairs we brought home. All laid between
1-7:00 a. m., July 4. Eggs smaller than those of Hyla femoralis. Do they
lay singly or below the surface? The outer envelope is very close to the
inner envelope." We finally concluded (1923, p. 406) they laid single eggs
deposited on the bottom. More observations are needed.

The actual count of the egg complement of one of the four females (above
mentioned) which laid its eggs in captivity was 942 . Another egg complement
count is 972.

320 Frogs of the Okefinokee Swamp

In 1922 field examinations of the eggs gave the following summation:
'Mtellus .8-1.0 mm.; inner envelope 1.2 -1.6 mm., outer envelope 1.4- 1.8 mm.
Eggs brown above, cream below."

Laboratory examinations gave .9 mm. as average vitellus, range .8-1 .0 mm.
inner envelope, average 1.4, range 1.4-1.6 mm.; outer envelope average 1.6
mm., range 1.4-2.0 mm.

Dangers. On August 11, 1922, at Camp Pinckney "we found countless
males in temporary pools 1-3 inches deep. Must be that they lay single eggs.
Must lay in shallow pools. No end of the tadpoles must be lost from drying
of breeding places."

HATCHING PERIOD

The eggs laid July 3-4 hatched in laboratory in about two days. In fact
some of the eggs (No. 998) laid July 4, i :oo-7 :oo a. m. hatched July 5 at
12:00 noon or 36-43 hours after ovulation or i 1/2 or i 3/4 days required for
liatching. This was in a period when maxima averaged near 88-90° and
minima close to 7o°F.

TADPOLE

Color description from life {June 23, 1922). General color of body citrine
drab. In general it has a greenish cast hke tadpoles of Hyla cinerea and H.
gratiosa. Developing ventral flap between hind legs ecru-olive; throat from
line of eye to eye to mouth same color. Area behind gill area ocher red.
Belly pigmented testaceous and chalcedony yellow. These make a solid
iridescence which on the sides and across the middle venter becomes replaced
with black.

Tail. Muscular part of tail dark olive buff. Muscular part without
prominent clusters of blackish spots. The dots are uniformly scattered over
the muscular part but slightly thicker near the tip. The tail reminds one
more of a Ranid tadpole tail in coloration.

Iris more or less black with a rim of testaceous or light coral red and
chalcedony yellow spots amongst the black.

General appearance. Tadpole small (32 mm.) full and fairly deep bodied.
Tail long, tail tip acuminate a fiagellum present. Dorsal and ventral crests
well developed and tail in general deep. Dorsal crest not so deep as the tail
musculature. Dorsal crest extends on to the body about to the vertical of
the spiracle. Spiracle sinistral directed upwards and backwards, well below
the lateral axis, the spiracular opening prominent elliptical lateral axis, is in
dorsal aspect on the lateral outline and in consequence is visible from the
venter. Anus dextral, opening at the level of the edge of the ventral crest.
Muciferous crypts indistinct.

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
upper fringe for about 1/2-3/10 of upper fringe. The end of the second upper
labial row is usually even with the end of the upper fringe. The horny beak
is contained about 1.8-2.0 times in the upper fringe. The median space
between the lateral second upper labial rows of teeth short, 3.25-5 in the length

Hyla squirella 321

of either lateral row sometimes very narrow (6 in lateral row). The inner
papillae extend under the third row of lower labial teeth making at least two
rows across the lower labial border. In the lower labial corner there is a
heavy papillary series sometimes 3 or 4 rows wide like H. versicolor or //.
femoralis. The lower third labial row is long and is usually about the length
of the first or second lower labial rows and longer than the horny beak. The
first and second rows are about equal and i 1/2-1 3/4 greater than the horny
beak. This species, like H. versicolor and H. femoralis has the upper fringe
very angulate at the middle.

Measurements. Length of body (10-12 mm.) in tail (16-20 mm.) 1.55-2.05,
average 1.75. Width (5.0-6.6 mm.) of body in its own length 1.7-2.2, average
1.875. Depth (5,0-7.0 mm.) of body equals body width, or slightly greater.
Depth of body 1.5-2.2 in body length, average 1.81. Depth (6.0-8.0 mm.) of
tail in length of tail 2.3-3.3, average 2.8. Muscular part (3.0-4.2 mm.) 1.5-
2.33 in depth of tail, average 1.87. Spiracle 1.15-1.75 nearer base of hind
legs or vent region (4.0-6.5 mm.) than the tip of the snout (6.5-8.0 mm.),
average 1.47. Spiracle 1.4-2.2 nearer eye (2.8-3.5 mm.) than base of hind
legs or vent, average 1.7. Eye 1.1-1.6 nearer to spiracle (2.8-3.5 mm.) than
to snout (4.0-5.0 mm.), average 1.38. Nostril 1.5-2.4 nearer eye (1.2-2.0 mm.)
than snout (2.8-3.5 mm.), average 1.8. Mouth (2.0-3.5 mm.) usually equal
to internasal space (2.5-3.5 nim.). Mouth contained 1.1-1.8 (average 1.50)
in interorbital distance (4.0-5.5 mm.). Internasal space contained in inter-
orbital space 1.33-2.0, average 1.58.

The dimensions of the largest tadpole are:

mm.

mm.

Total length

32 ■

Spiracle to vent

6.5

Body length

12

Spiracle to eye

3-0

Body depth

6.5

Eye to snout

4.0

Body width

6.0

Eye to nostril .

1 . 2

Tail length

20.0

Nostril to snout

3-0

Tail depth

6.0

Mouth

3-2

Musculature of tail

4.0

Interorbital distance

5-0

Spiracle to snout

7-5

Internasal distance

3-5

General remarks. In 1923 (p. 406) we characterized these outstanding
characters as "bodies greenish, tail uniformly sprinkled with black dots."

LARVAL PERIOD

On July 3, 1922, at Trader's Hill, we found the species very active in
congress. On August 9, ^^ days later there were "small tadpoles at Trader's
Hill ponds. If the tadpoles transforming July 10 ,were laid in the very rainy
last week of May, 1922, they were 40-45 days in the larval life.

The eggs laid in camp July 3-4 hatched July 5 developed by July 30 to
tadpoles 1 1 mm. long but proper food conditions were not at hand. Some of
them fixed August 21 measured 16 mm, or one-half normal tadpole size 48

322 Frogs of the Okefinokee Swamp

days after ovulation. Even with these 25 or 30 days more or 50-55 days
probably would have sent them to transformation if put into natural pond.
40-50 days or even two months may elapse for the larval period.

TRANSFORMATION

Period. On June 26, 1922 we "found our first series from tadpole to
transformation in Anna's Pond 3/4 mile northeast of Starling Branch crossing.
It is an open pond or sink hole with no trees nearby." On the following day
we made more records of tadpoles. On July 10 the Hyla squirella tadpoles
at above pond were much fewer. "Some of them transformed and some of
them have left the pond." Our transformation notes of actual transformation
are from June 2 6- July 11 but eggs laid in July and August 1922 must trans-
form much later.

Size. One lot taken June 26, 1922, measured 11, 11, 12, 12, 12, 12, 12, 12,
12.5 and 13 mm. respectively. Seven taken June 27 measured 11.5, 12, 12,
12, 12, 12 and 12.5 mm. respectively. Another lot of twenty specimens had
one at 11 mm., two at 11. 5, eleven at 12 mm., four at 12.5 mm., and two at
13 mm. One group of six collected July 11, 1922 have the following points of
interest: i mature tadpole; one 12 mm. long, tail 12 mm., forelimbs almost
ready to burst through; complete tadpole mouth; a third 12 mm. long, tail
15 mm., tail filament, forelegs through, tadpole mouth but without teeth; a
fourth 12.5 mm., tail 13.5 mm., tadpole mouth just past; a fifth 11 mm., tail
10 mm., forelimbs not quite out, tadpole mouth complete except for third
lower row of teeth; and a sixth, 11.5 mm., tail 11 mm., no tadpole mouth,
mouth half frog-like, fore legs out. The range of the forty-one specimens is
from 11-13 mm., the mode 12 mm., the average 12.3 mm.

General remarks. These httle frogs are best identified by their tails as
long as they have the crests. Thereafter no key could identify them. They
have not the whole line on upper lip, no spots on back, nor vitta back of eye.
Like many Hyla transformed frogs they are hard to distinguish unless followed
through the stages or unless tadpoles be known.

GROWTH

On June 26 and 27 we had 36 specimens 11- 13 mm. at transformation.
On August 16, 1910, Professor J. C. Bradley took one at Lakeland, Fla., 14
mm. in length. Seven miles north of Millen, Georgia, we took June 10, 1922,
two Hyla squirella of two growth groups one 20 mm., and a male 28 mm. All
the twelve croaking males of July 3, 1922, and all the four pairs of July 3,
1922 (another pond) are from 27-35 mm. in length. Have we three forms
11-14 mm. at transformation; 20 mm. one year old and 27-35 two year old?

In 1922 (March 9) Hurter took (U. S. National Museum Nos. 581 17-
58124) specimens of two groups, two 22 and 23 mm., and six 27.5, 28, 29.5,
3*2, 33, 33 mm. LeConte in Liberty County took two (U. S. National Museum
No. 3645) 21 and 27 mm. respectively. Dr. Schwartz and Mr. Barber (U. S.
N. M. 61690-1) at Paradise Key, Fla., took two 21 and 26 mm. On July 18,

Hyla squirella 323

1901, P. R. Paine at Charleston, S. C, took 16 specimens (U. S. National
Museum No. 29229-29243) 17.5, 17.5, 19, 19.5, 20, 20, 20, 20.5, 21, 22, 22, 23,
23, 23, 24.5, 25 mm., respectively 17.5-25 mm. William Wittfield took (U. S.
National Museum No. 12008) twelve specimens 15, 16, 19, 21, 24, 24, 25, 25,
26.5, 27.5, 28, 28.5 mm. respectively. Material taken by Professor A. M.
Reese, Lake Kissimee ranges from 20-27 mm. (one 31 mm.) for 23 specimens,
(U. S. N. M. Nos. 36202-10, 29065-66, 36249-36260), an average of 23.5 mm.
All combined it seems as if 11-14 mm. is transformation size, 17.5-25.5 mm.
is one year old size, 25-35 mm. is two year old size.

Very recently we discovered considerable material in the U. S. National
Museum from one locality, Charleston, S. C. in the fall of 1923 and the
summer and fall of 1924. They were taken by J. T. Rogers as follows. On
June 19, 1924, he secured (U. S. N. M. Nos. 67425-42) all adults 23-33 mm-
(Males 24, 25, 25, 27, 27, 28, 28, 33; females 23, 24, 24, 25, 25, 25, 26, 26, 30)
— all possible two year olds 23-33 i^i^- but the 30 and 33 mm. individuals are
possibly apart. On Sept. 17, 1924, J. T. Rogers took (U. S. N. M. Nos. 67714-
67730) the following: 14.5, and 17, 18, 18, 18, 19, and 21, 22, and 26.5, 27, 27,
27, 28, 28, 28, 29, and 32 mm. Have we three groups, 14.5 mm., 17-22 mm.,
26.5-32 mm.? On November 16, 1923, Rogers took 4 specimens (U. S. N. M.
66804-7) i7-5j 20, 20, 22 mm. Does all this material shape as follows: 14.5
mm.; 17-22 mm.; 23-30 mm.; 32, 33 mm.?

After measuring the American Museum material their data seems to fall
thus: 13 mm. transformation; 19-23 mm. or 25 mm. for the first year olds; 25
or 26-36 mm. for 2 year olds. Several specimens from 19 to 23 mm. or in one
instance 25 mm. we recorded as doubtful in sex. Doubtless they were all one
year olds. But the males and females clearly so externally 23-25 mm. are
doubtless individuals older than one year.

FOOD

Deckert (192 1, p. 22) in Miami, Dade Co., Florida, finds that "A few have
established themselves in the velvet bean vines covering the writer's back
porch, and at night will sit flattened out against the window panes, catching
the insects that are attracted by the light within."

ENEMIES

On June 27, 1922, in one pond "we found a Hyla squirella tadpole with
one hind leg. Yesterday found one tadpole with forelegs about to burst out
but it had no hind legs. Either lost or never developed." In this pond we
took southern riband snakes and chicken turtles {Deirochelys).

AUTUMNAL DISAPPEARANCE

Deckert (192 1, p. 22) records them after heavy rains in September in
Dade County, Florida. Pope (19 19, p. 97) at Houston, Texas, heard it as
late as September 20. "Have heard very httle of Hyla squirella lately, but
rain fell all last night and tonight they were singing loudly again, although it

324 Frogs of the Okefinokee Swamp

turned cool after the rain." Our latest record is September 17, 1922, (F.
Harper). In the U. S. National Museum are the following captures: J. T.
Rogers, Sept. 17, 1924, Charleston, S. C. (U. S. N. M. 67714-67730); same
collector, Nov. 16, 1923 (U. S. N. M. 66804-7); J- T. Rogers, Sept. 15, 1924
(U. S.N. M. 67731).

AFFINITIES

In 1855 (p. 429) LeConte holds "It is wrong in Dumeril and Bibron to
say that this species {Hyla femoralis) is a variety of Hyla squirella. In shape
and size the difference is most considerable. The latter animal during the
warm season is always to be met with about houses, the H. femoralis never.
Besides, their notes are entirely different."

Cope (1889, p. 364) maintains that "This species approaches nearly some
forms of H. regilla, of which it may be considered the southeastern representa-
tive, though the regions inhabited by both approach in Texas. It may be
always distinguished by the more depressed head and weaker canthus ros-
tralis; it is rarely so robust and usually of more delicate and less striped colora-
tion. H. miotympanum Cope replaces it in Mexico; the resemblance between
them is offset by greater palmation and smaller tympanum of the latter."

Hyla squirella on the basis of eggs, if judged on manner of deposition,
would fall in the group of singly deposited eggs: Hyla gratiosa, Hyla andersonii
Acris gryllus, Hyla crucifer, but this may mean little. But if we analyse the
egg characters it may fall closer to Hyla femoralis, Hyla cinerea, etc. In our
synopsis of tadpoles, Hyla squirella apparently falls with Hyla versicolor, Hyla
arenicolor, Hyla femoralis, the group with long lower hind labial row. On
June 17, 1922, we note that "the tail of the tadpole reminds us of the tail of
some Ranids in color."

BIBLIOGRAPHY

1907 Brimley, C. S. Jour. Elisha Mitchell Soc. Dec. 1907. Vol. XXIII, No. 4, p. 154.
1889 Cope, E. D. U. S. National Museum Bull. 34, 1889, p. 364.

1914 Deckert, Richard F. Copeia, Feb. 14, 1914, No. 3, p. 3.

1915 . Copeia, May 15, 1915, No. 18, p. 3.

1921 . Copeia, March 15, 1921, No. 92, p. 22.

1922 . Copeia, Nov. 20, 1922, No. 122, p. 88.

1842 De Kay, James E. Zool. of N. Y. Part III, p. 72.

1903 Geyer, Hans. Natur und Haus Jahrgang XI, Heft 7, Jan. i, 1903, pp. 97-99.

1842 Holbrook, John Edwards. North American Herpetology. Vol. IV, Phila., pp.

123-125.

1924 Holt, E. G. Copeia, Nov. 18, 1924, p. 95.

1825 LeConte, John. Ann. Lye. Nat. Hist. N. Y., Vol. I, Part II, N. Y., 1825, pp. 38, 39.

1855 . Phila. Acad. Nat. Sci. Proc. 1855, P- 429-

1924 Myers, George S. Copeia, June 30, 1924, No. 131, p. 60.

1919 Pope, Phihp H. Copeia, Dec. 31, 1919, No. 76, pp. 96, 97.

1924 Schmidt, Karl P. Copeia, July 15, 1924, No. 132, p. 68.

1923 Viosca, Percy J. Copeia, Feb. i, 1923, No. 115, pp. 38-40.

1923 Wright, A. H. The Anatomical Record, Jan. 20, 1923, Vol. 24, No. 6, p. 406.

1923 • Copeia, Feb. i, 1923, No. 115, p. 34.

1924 and A. A. Wright. The American Naturalist, July, August, 1924, Vol.

LVIII, p. 378.

Hyla versicolor 325

Hyla versicolor (Le Conte)

(PI. II, Fig. 9; IV, Figs, 3, 8; VII, Fig. 2; X, Figs. 9, 10; XII, Fig. 12; XV, Fig. 6; XVII;

XXXIII; Text Figs, i, 18)

{See Life History in Carnegie Publication No. 197, 1914, PP- 44-51).

COMMON NAMES

Common Tree Toad. Common Tree Frog. Tree Toad. The Northern
Tree-toad. Chameleon Hyla.

RANGE

Check list. Maine, southern Canada, west to Minnesota, south to the
Gulf States (Texas and Arkansas in part only). — Stejneger & Barbour, 1923,
p. 32.

Supplementary records. Edith R. Force (1925, p. 25) records Hyla versi-
color in Okmulgee County, Oklahoma, in 1924.

In Alabama Holt (1924, p. 95) records this species from Sand Mountain,
Jackson County.

In 1 91 7 we found it on May 31 at Petersburg, Va.; on June 2 at Din-
widdle, Va.; on June 11 several near Flatwood, Ala. From 1913 to 1925, we
have taken this species from Dorset, Ontario, to near Jacksonville, Fla., and
thence west to Beeville, Texas.

Local Okefinokee records. In 192 1, F. Harper recorded this species about
ten miles south of Moniac on the Georgia side. Again "during the evening
of August 16, after a thunderstorm several individuals were found in voice at
Camp Pinckney." In 1922 we heard them from June 16 onward at Camp
Pinckney, Trader's Hill, Spanish Creek Woods, Thompson's Landing, St.
Mary's River, also along road from Folkston to Chesser Island.

MEASUREMENTS

(Recent Material)

Head to angle of mouth i.o (18 mm. transformation)-!. 33 (28 mm.)-
1.41 (44 mm. c?')-i.5 (56 mm. 9) in length; head to rear of tympanum
(absent)-i.2 (28 mm.)-i.i7 (44 mm.)-i.i7 (56 mm.) in length; head to
angle of mouth 3.6-3. 1-3.66-4. i in length of body; head to rear of tympanum
( )-2.8-3.o-3.3 in length of body; snout .66-.63.1.07-.94 in first finger;
snout 1.0-1.1-1.15-1.3 in fourth finger; snout .5-.54-.7-.60 in first toe; eye
1. 2-1. 37-1. 08-1. 4 in snout; eye ( )-.75-.5o-.66 in tympanum; eye .8-.87-
1.16-1.33 in first finger; tympanum ( )-i. 83-2. 16-2. 12 in snout; tympanum
( )-3. 0-4. 5-4.0 in intertympanic width; internasal width .75-.83-1.12-1.0
in upper eyelid space; interorbital width .8-1.2-.9-.83 in internasal width;
interorbital width .60-1.0-1.0-.82 in upper eyelid width; interorbital width
( )-3. 6-3. 1-2.66 in intertympanic width.

Forelimh: Forelimb i. 8-1. 4-1. 9-1. 6 in length of body; forehmb 2.2-2.2-
2.5-2.12 in hind limb; first finger 2. 0-2. 3-1. 3-1. 6 in hind finger; second finger
1. 6-1. 6-1. 2-1. 3 in third finger; second finger .8-.7-.94-.8 in first finger; third

326 Frogs of the Okefinokee Swamp

finger .62-.62-.83-.77 in second finger; fourth finger i. 33-1. 25-1. 2-1. 18 in
third finger; fourth finger .5-.5-.6-.45 in first toe; internasal width 1.0-1.16-
1. 75-1. 6 in first finger; internasal width i. 25-1. 6-1. 87-2.0 in second finger;
internasal width 2.0-2.3-2.25-2.6 in third finger; internasal width 1.5-2.0-
1,87-2.2 in fourth finger.

Hindlimh: length i. 22-1. 5-1. 36-1. 25 in hindhmb; tibia 2.5-1.7-2.2-2. 15
in length of body; tibia 3.1-2.75-2.95-2.7 in hindlimb; tibia i. 4-1. 25-1. 15-
1.26 in forehmb; tibia .71-.875-.90-.92 in hind foot; first toe i. 66-1. 5-1. 33-
1.8 in second toe; first toe 2.33-2.66-2.2-3.33 in third toe; first toe 3.00-3.66-
2.9-3.9 in fourth toe; first toe 2.33-2.66-2.1-2.4 in fifth toe; second toe 1.8-
1. 8-1. 66-1. 77 in third toe; second toe 1.8-2.44-2. 16-2. 16 in fourth toe;
second toe i. 8-1. 8-1. 6-1. 33 in fifth toe; third toe i. 3-1. 3 7 5-1. 3-1. 2 in fourth
toe; third toe 1.0-1.0-.93-.75 in fifth toe; fourth toe 1.1-1.27-1.4-1.23 in
hind foot; fourth toe i. 55-1. 45-1. 54-1. 3 in tibia; fourth toe 2. 2-1. 8-1. 77-1. 7
in forelimb; fifth toe i. 3-1. 3 7 5-1. 3 5- 1.6 in fourth toe; internasal width .75-
i.o-i. 22-1.0 in first toe; internasal width i. 25-1. 5-1. 5-1. 8 in second toe;
internasal width 1.75-2.66-2.5-3.33 in third toe; internasal width 2.25-3.66-
3.25-3.9 in fourth toe; internasal width 1.75-2.66-2.4-2.4 in fifth toe.

HABITAT

This form we found only from ten miles south of Moniac on the St. Mary's
River to Folkston really Camp Pinckney on the St. Mary's River but not
within the swamp.

Holbrook (1842, Vol. 4, p. 117) writes "This animal is commonly found on
trees and about old stone fences, overgrown with mosses and lichens, the color
of which it so closely resembles that it frequently escapes observation even
when sought for. It very commonly chooses old and decaying plum trees for
its abode, probably because insects on which it feeds are most abundant in
such situations."

FIRST APPEARANCE.

In Ithaca our first appearance records for this species come in mid April
or earlier. Surely in Southern Georgia they must be abroad even earlier in
the spring. In 192 1 we spent April and May within the swamp where the
species is not but in 1922 we encamped in June and on June 19 we heard them
in Camp Pinckney.

In Houston, Texas, Pope (1924, p. 95) made "the following notes on it:
Feb. 13, 19 18, Have heard several in the past few days calling from trees in
camp, but have not seen any yet. April 15 collected one specimen, a male
that was calling from the branch of a pine tree. April 24 found one calling
in rain pool where H. squirella was breeding. I heard them frequently on
warm evenings, answering each other from trees in the woods near camp.
After the first of May they became silent and I heard and saw no more of them
for the season. If they laid eggs in any of the pools near camp they made no
such noise about it as they do in the North." We heard and collected them
in Beeville, Texas, March 24, 1925.

Plate XXXIII
Tree-toad {Hyla versicolor)

1 . Old turpentine pit at Camp Pinckney, Ga. June 22, 1^22.

2. Male croaking, Camp Pinckney, Ga. August 11, 1922. Flashlight.

3. Male, Beeville, Texas. March 26, 1925. Dorsal aspect, x 0.8.

4. Male croaking on pine tree about i^ feet above ground, Camp Pinckney, Ga.

August 16, 1 92 1. Flashlight.

5. Transformed frog, Chesser Id., Ga. June 29, 1922. x i.o.

6. Egg packet attached to Potamogeton leaf, Ithaca, N. Y.

7. Female and male, Chesser Id., Ga. August 12, 1922. Ventral aspect, x 0.60.

Hyla versicolor 327

GENERAL HABITS

Metachrosis. LeConte (1825, p. 281) called it Hyla versicolor in allusion
to its metachrosis. Harlan (1927, p. 343) in his description alluded to its
"colour varying with the will of the animal from pale-brown to cinerous and
green." Holbrook (1842, Vol. IV, p. 115) spoke of it thus: "This is a beautiful
species of Hyla: its colours seeming almost to vary at the will of the animal."
DeKay (1842, Part III, p. 71) says "its facility of assimilating the color with
that of the tree on which it rests, renders its capture very difficult." Since
that date writers have alluded to this phase of the tree toad's life more than
any other one topic. It is too well known to require further discussion.

VOICE

This is a famihar phase of its life history. Holbrook (1842, Vol. IV, p. 117)
writes :

"It is very noisy towards evening in cloudy weather or before rain, its
voice consisting of a liquid note, terminating abruptly, like 1-1-1-1-l-l-luk. At
the close of spring, and during a great part of the summer, when the toad has
become silent, this note may be heard especially in the evening from various
shallow pools, to which the animal resorts for the purpose of depositing its
spawn."

DeKay (1842, Part III, p. 71) asserts "I have been assured by many
credible persons that it possesses ventriloquial powers in no considerable
degree, and often deceives the most attentive observers."

Cope (1889, p. 375) characterizes it as follows: "Its voice is a loud, coarse,
resonant trill, uttered with a uniform pitch, and continued for two or three
seconds. It is heard about bodies of water in the spring, when the sexes are
depositing and fertihzing the eggs. Later in the season it proceeds from fences,
hedgerows, and orchards, as well as from the forest, often at no great elevation
from the ground. They are especially noisy towards evening after a rain."

In 1906 Fowler says that "The vocal sac when inflated is very marked,
though apparently it does not distend to the extent of that of pickeringii.
It collapses by a series of jerks as the cry is emitted. They usually do not
begin to call much before late April."

The same year Dickerson (1906, p. 119) writes: "At dusk or on rainy days
a loud resonant trill comes from the trees and vines. The sound has the charm
of contentment in it; in fact it is much like the purring of a cat, only louder.
At a distance it sound something like the bleating of a lamb. The pitch is
uniform, but may vary with the individual, from G above middle C to E
above. It continues for two or three seconds at a time, then ends abruptly.
It may be given several times in rather close succession. If we see the tree
frog trilling, we are surprised that the whole body is so greatly agitated and
that the throat extends into so large a sac. The size of the sac decreases be-
tween each two trills and at the end collapses, leaving a very wrinkled throat."

Overton, a pioneer in amphibian flashlight photography, has studied the
caUs of our northeastern frogs very assiduously. He discusses this form thus.

328 Frogs of the Okefinokee Swamp

"The voice of the tree frog is a loud musical trill, like a low-pitched whistle.
It may be recognized by its extremely pronounced trill. Each individual call
lasts about two seconds and may be repeated at intervals of ten to twenty
seconds.

"When a tree frog sings, it distends a throat pouch to about the size of its
head. The vocal pouch vibrates with the trilling and so its photograph will
often be blurred. The frog usually sings while sitting upright in a dry place,
such as a lily pad or a branch over the water. It is not especially shy, and is
usually easy to photograph.

"Besides the trill, the tree frog has another note that is not generally
known and recognized. This note is exactly like the sound made by a hen
turkey that is calling to her mates. It is a low, mournful "chow, chow, chow,
chow, chow," whistled with a descending pitch, repeated about three times a
second for two or three seconds. When I first heard the sounds, their source
was a mystery, for they were infrequent. But at last I discovered the musi-
cians. Sitting on a small limb just above the water were two tree frogs about
six inches apart. Turning about and facing each other, one gave its turkey
call by chirping, while nodding its head and body with every chirp-like and
puppy barking. It was at once answered by the other in the same way. The
two then quickly turned about and resumed their ordinary songs. This
performance was repeated several times while I watched them with aching
feet, for they waited a long time between their turkey calls. The Long
Island negroes name the call a 'turkey root' from a superstitious behef on
the virtues of roots in conjuring and magic. A negro calls any mysterious
sound or sight a conjure or "root,' and he will run a mile to avoid hearing a
'turkey root.' "

We (1914, pp. 45, 46) have discussed their voice and calling at length and
will not attempt its characterization here.

In 192 1 on my way to the Okefinokee Swamp I stopped at Raleigh, N. C.
where I heard males calling April 15 and many calling April 16.

In 1 92 1, August 16, at Camp Pinckney, F. Harper and Marion Lee of our
party heard this species. The former notes that: "Their habitat here was
about some temporary and permanent pools in level woods of live oaks and
other oaks, with a few pines. Some were trilUng on the ground, while others
in voice were clinging in a vertical position on the trunks of pine or oak within
a couple feet from the ground. The note is a rather vigorous high-pitched
trill: r-r-r-r-r-r-r-r-r-r-r-r-r-t, etc. Ten or eleven calls were given in the space
of half a minute and each one lasted about 2/3 of a second. At its maximum
inflation the throat sac attains a size a little larger than the head, and in the
intervals between trills it is deflated to about half that size."

In 1922 we heard it from June 19- August 12. On August 11 "at Camp
Pinckney from 1-3 p.m. were heard several males in the trees. At 8:30-10
p.m. we found several in the oaks. One in black gum (small tree 4 feet high)
beside pond No. 4 where we found Hyla versicolor tadpoles the last time we
came. Several males low in bushes on south side of Pond No. 3. Several
were in trees 10-15 feet up or more. Took a female on the bole of a tree near

June

19.

June

27.

July

9-

July

24,

August

9'

August

II.

August

13.

Hyla versicolor 329

Pond No. 5. She was perched side wise on the trunk and 3 feet or more farther
up was a male. In Pond No. 5 took an adult Hyla versicolor. These males
seem more marked and barred on hind legs than our northern tree toad and
more orange appears on the groin and hind legs."

On August 13 tardy breeders in Spanish Creek woods were calling loudly
and almost in chorus.

On our way from Ithaca to Okefinokee we heard it from Pennsylvania to
Georgia June 2- June 12, 1922.

A resume of our few records of calling in 192 1 and 1922 follows:

1921
August 8. 10 miles south of Moniac. Heard this species.
August 16. Several calling at Camp Pinckney.

1922
Several calling during the rain.
Several calling when rain threatened.
Thompson's Landing. Heard in the woods.
Heard in Spanish Creek Woods.
Heard near Trader's Hill two H. versicolor.
Several calling 1-3, more 8:30-10 p. m.
Several calling along Folkston-Chesser Road-Spanish Creek Woods.

At the nearby U. S. Weather Bureau Stations the temperatures were
maxima 8i°-96°, average 91°, minima 6 2 "-7 6°, average 70°. None of our
temperatures from 6:00 a.m. to 9:00 p.m. or sometimes midnight were below
71° for these periods of calling. As every one knows it calls most frequently
before, during and after rains.

MATING

Female. "Under parts entirely white, whiter than in the males. Other-
wise the same."

Structural differences. The males average slightly smaller. In several
pairs taken we have these measurements: Tallulah Falls, Ga., June 22, 1909,
cf 44 mm., 9 45 mm.; Camp Pinckney, Ga., Aug. 12, 1922, cf 45 mm.,
9 53 mm.; Ithaca, N. Y., June i, 1906, 6^ 46 mm., 9 50 mm.; Ithaca, N. Y.,
June 14, 1907, cf 46 mm., 9 50 mm.; Ithaca, N. Y., June 10, 1907, cf 48
mm., 9 59 mm.

The males have the dark vocal sacs. The underparts of the female are
pure white, rarely so in the male. There is little of difference in fingers or
tympana in the two sexes. Probably two-year-olds begin breeding. A male
30 mm. has a dark throat.

Duration. Mating takes place largely at night and a pair mated at night
usually lays the same evening as mated.

Amplexation. In this species the embrace is axillary. On August 11,
1922, 8 :3o-io:oo p.m. "at Camp Pinckney we took several males. Two males
were put together, embraced and continued thus for several hours. It was an
axillary amplexation. In this pair the upper male was much lighter, the lower
one much darker."

330 Frogs of the Okefinokee Swamp

OVULATION

Habitat. We made little effort for the life history of this species in
Okefinokee Swamp country. We found their tadpoles in water pits at Camp
Pinckney; and in suitable ponds and places near Spanish Creek Woods they
must have laid. On June i, 191 7, when we did not know Acris eggs, at Din-
widdie, Va., we found in a shallow pool (10 x 8 ft. in diameter and 4 inches
deep) no end of eggs packets on the surface. Normally we would have in-
stantly called them Hyla versicolor but we had heard Acris in this direction.
When nightfall came, in the bushes around this pool were several Hyla
versicolor males. They were Hyla versicolor eggs not those of Acris.

Period. At Ithaca we have records of breeding from May 10- June 17.
They were active in 1922 from June 3- June 12 onward on our trip from Ithaca
to Georgia. We heard them several times in Pennsylvania, at New Alexandria,
Va., Roanoke River, Cary, and Everett's Pond, N. C. We heard them in the
Okefinokee country from June 10- August 13, 1922. We might not be inclined
to think of them breeding, were it not that a female taken August 1 1 had ripe
ova. Mature tadpoles (some with two legs) June 22, must have been laid in
early days of May, as early as May i, or the last of April. On July 5, 1922,
we found eggs at Camp Pinckney, Ga., eggs which we suspected were those of
Hyla versicolor.

Temperature and humidity. The eggs we found July 5, 1922, were laid in
the period of rain (every day from July 1-5) when 2.97-3.20 inches fell, when
minima reached from 68°-72°, and maxima from 83°-96°. The greatest
period of activity came August 8-13 when minima were 68°-74°, maxima
8i°-93° and rainfall for this period was 2.35 inches, 1.48 inches, 1.49 inches,
2.35 inches and 4.06 inches for stations around the swamp.

Egg-laying process. In 19 14 (pp. 48, 49) we described the process,

EGGS
In Dinwiddle, Va., June i, 19 17, the tree toad eggs were as recorded for
Ithaca, N. Y. Some of the packets were larger than ever recorded before.
Occasionally it looked as if several packets were merged. In 1924 (p. 378) we
characterized the eggs as follows: "Eggs deposited in a mass. Egg mass a
surface film. Egg envelope outhne indistinct, more or less merged in the jelly
mass; jelly gelatinous; egg brown above, cream or yellow below. Egg packets
small, masses seldom if ever over 20 sq. in. (125 square centimeters), or 4 by
5 inches in diameter (10 by 12.5 cm.). Inner envelope small 1.4 to 2.0 mm.;
outer envelope 4 to 8 mm. Packets small, seldom over 30 to 40 eggs; vitelline
I.I to 1.2 mm. Egg complement, 1802. Season May 10 to August 12."
The above complement is compiled from a female captured August 12, 1922.
The count of the left ovary was 901.

TADPOLE

Color description from life ( Non Ridgway) . General color of back olive
green. Background of back yellowish, several with many fine hair-like black
markings and golden and black spots, becoming orange in the head region

Hyla versicolor 331

and sometimes almost vermillion about the eye; the golden and black spots
are more pronounced toward the tail; on the sides is a decided iridescence.
The eye is slightly bronzy. The venter in general is conspicuously white or
light cream and slightly iridescent. The belly is covered with fine golden
spots. From the gill-region forward the venter is greenish, a coloration pro-
duced by black and golden spots. The background of all the tail except the
base is scarlet or orange- vermillion. The base of the tail is like the body.
The tail is covered with black blotches, more prominent around the edges of
the crests. These blotches become much more numerous as the hind legs
develop.

General appearance. Tadpole medium (46.6 mm.) full and deep bodied.
Tail very long, deep, tip very acuminate with a prominent fiagellum. The
dorsal crest is as deep as the musculature, and extends on to the body to the
vertical between the spiracle and the eye or to the spiracle. Spiracle sinistral,
directed more backwards than upwards, far below the lateral axis and visible
as an elliptical opening. Eye on lateral axis, in dorsal aspect on the lateral
outline and in consequence visible from the venter. Anus dextral, very near
or at the level of the edge of the lower tail crest. Muciferous crypts indistinct.

Mouth parts: Teeth 2/3. Upper labium fringed with a continuous row of
labial teeth; the papillae extend above and inwards beyond the end of the
upper fringe for about 1/3 of the length of the upper fringe. The end of the
second row usually is even with the end of the upper fringe. The horny beak
is contained about 2 times in the upper fringe. The median space between the
lateral second upper labial rows quite short, 3.25-5 times the length of either
lateral row. The inner papillae extends under the third row of lower labial
teeth making at least two rows of papillae across the lower labial border. In
the lower labial corner there is a heavy papillary series of 2 or 3 rows not so
pronounced as in Hyla squirella or Hyla femoraUs. The lower third labial
teeth is long and is contained usually about 1.10-1.25 times in the first or
second row. The first or second rows about equal and 1.8-2.0 greater than
the horny beak. This species like Hyla femoralis and H. squirella has an
angulate upper fringe at its middle.

Measurements {Okefinokee specimens). Length of body (14. 5-1 6. 5 mm.)
in tail (27.4-21.2 mm.) 1.75-2. i average 1.22. Width (9.6-1 1.4 mm.) of
body in its own length i. 38-1. 6 average 1.5. Depth (8.2-10.0 mm.) of body
1. 0-1.2 in body width, average 1.08, Depth of body 1.5-1.8 in body length,
average 1.64. Depth (6.6-9.0 mm.) of tail in length of tail 3.1-4.0, average
3.6. Muscular part (4.6-5.0 mm.) i. 72-1. 9 in depth of tail, average 1.6.
Spiracle 1.3-1.6 nearer base of hind legs or vent region (6.6-9.0 mm.) than the
tip of the snout (9.8-1 1.4 mm.). Spiracle i.i 2-1.5 nearer eye than base of
hind legs or vent, average 1.33. Eye to spiracle (5.0-6.0) about equal to eye
(5.0-6.0 mm.). Nostril i. 36-1. 6 nearer eye than snout. Mouth (3.4-4.0
mm.) 1.1-1.36 in internasal space (4.2-4.8 mm.), average 1.25. Mouth con-
tained 1.65-2.05 (average 1.8) in interorbital space 1.35-1.6, average 1.45.

332 Frogs of the Okefinokee Swamp

The dimensions of the largest tadpole are:

mm.

mm,

Total length

46.6

Spiracle to vent

7-6

Body length

15-8

Spiracle to eye

6.0

Body depth

10. 0

Eye to snout

5-0

Body width

II. 4

Eye to nostril

2 . 2

Tail length

30.8

Nostril to snout

30

Tail depth

6.8

Mouth

3-6

Musculature of tail

5-0

Interorbital distance

4.2

Spiracle to snout

10. 0

Internasal distance

6.0

These tadpoles have long tails, 1.75-3.5 in the length of the body and
scarlet or vermillion in color, with black blotches more prominent near the
margins of the crests. The intenstine does not show through the skin.

LARVAL PERIOD
In the North it takes 50 or 60 days for development from the egg to trans-
formation. Sometimes the period may be only 45 days or in other cases as
great as 65 days. In the Okefinokee country the period may be shorter.

TRANSFORMATION

Period. The only transformation we found came June 27, 1922 at Camp
Pinckney. "In the Hyla versicolor pond some are almost transformed."
This is almost a month eariier than any we have recorded at Ithaca, N. Y.
(July 1 8- August). Were they (June 27, 1922) tadpoles from eggs laid in the
first (May 1-5), middle (May 15-22), or last of May (May 28-31) when rain
prevailed, either 57 or 52 days or 42-36 days or 33-28 days?

Size. In 1914 the size of 87 specimens at Ithaca measured 13.6-20 mm.
at transformation, the average 1.6 mm., the mode 16 mm. The material
transforming June 27, 1922, at Camp Pinkney measures as low as 13 mm. in
body size.

GROWTH

In 1920, we concluded that tree toads were 14-20 mm. at transformation,
average 16 mm.; at i year old 20 mm.-3o mm., average 25 mm.; at 2 year old
30-41 mm., average 35 mm.; at 2 year old 41-51 mm., average 45 mm. If
they transform at 14-20 mm. the specimen we found at Ithaca, May 16, 19 15,
must have been transformed at the smaller size of 14 mm. It measured 18.5
mm. and was doubtless of the previous fall's transformation. The smallest
breeding frog (outward signs) was a male taken at Petersburg, Va., June i,
1 91 7. The males taken the same day at Dinwiddle were 34, 35, 37, 38, 39,
40, 41, 42 mm. The frogs taken at Camp Pinckney August 16, 1921, June 16,
1922, and August 11, 12, 1922 measure 39, 40, 41, 42, 42, 43, 43, 44, 44, 45 all
males and one female 53 mm. Transformation came at 14-20 mm. Possibly
20-30 mm. represents one year olds; 30-42 mm. represents two year olds;
42-53 mm. three year olds; 53-59 mm. four year olds. The largest we have is
a female 59 mm. in length.

Hijla versicolor 333

AUTUMNAL DISAPPEARANCE

Our latest record at Ithaca, N. Y., is October 24, 1905. The latest record
we have for the Okefinokee country is Aug. 12, 1922. It must continue to be
abroad until November at least. Harlan (1835, p. 109) relates how "A
specimen was dug up from the root of an apple tree in Bartram's Botanic
Garden, in the winter of 1828, several feet beneath the surface."

AFFINITIES

In 1825 (1825, p. 282), LeConte writes: "It is surprising that this species,
so common in the part of America that has been most visited by foreign
naturalists, has as yet been noticed by no one. It may have been confounded
with the H. viridis of Europe, which, however, it but little resembles."

In 1842 Holbrook (1842, Vol. IV, p. 117) alludes to the same oversight.
"It is remarkable that an animal so common and so very noisy should so
long have escaped the attention of naturalists. The first mention made of it
is in Kalm's 'Travels in North America' ; he, however, only describes its habits,
and refers to Rana arborea of Linnaeus, to which it bears but a slight resem-
blance "

LeConte (1855, p. 428) writes that "This animal is the analogue of the
Hyla arborea of Europe."

On method of egg deposition it lays packets on the surface like Hyla
femorahs. On egg characters it seems to fall with Hyla femoralis and Hyla
arenicolor, but the latter lays single eggs. On coloration of tadpole tail Hyla
versicolor is more like Hyla arenicolor than Hyla femoralis, yet all three may
have scarlet or vermilion in the tail. On tadpole mouth characters it falls
into the group with long lower third row of labial teeth. The adult reminds
one more of Hyla femoralis or Hyla arenicolor than other Hylas of the LTnited
States.

BIBLIOGRAPHY

1889 Cope, E. D. U. S. Nat. Mus. Bull. No. 34, p. 375.

1842 DeKay, James E. Zool. of N. Y.. Part III, p. 71.

1925 Force, Edith R. Copeia, April 30, 1925, No. 141, p. 25.

1906 Fowler, H. W. N. J. S. Report 1906, p. 114.

1827 Harlan, R. Journ. Acad. Nat. Sciences Phila., Vol. V, Part III, 1827, p. 343.

1835 . Medical and Physical Researches Phila., 183^, p. 109.

1842 Holbrook, E. N. A. Herpetology Vol. IV, p. 117.

1924 Holt, E. G. Copeia, Nov. 18, 1924, No. 135, p. 95.

1825 LeConte, John. Ann. Lye. Nat. Hist. N. Y., Vol. I, Part II, 1825, p. 282.

1855 . Phila. Acad. Nat. Sci. Proc, 1855, p. 428.

1914 Overton, F. Mus. Brooklyn Inst. Arts and Sciences, Science Bull. Vol. II, No. 3,

pp. 32, 33. Nov. 3, 1914.

1919 Pope, P. H. Copeia, Dec. 31, 1919, No. 76, p. 95.

1914 Wright, A. H. Carnegie Publication No. 197, Sept. 8, 1914, pp. 48, 49.

334 Frogs of the Okefinokee Siuamp

Rana aesopus (Cope)

(PI. Ill, Fig. 5; IX, Fig. 4; XI, Figs. I, 2; XIII, Fig. 4; XVI, Fig. 2; XVII; XXXIV; XXXV;

Text Figs. I, 22)

COMMON NAMES
Gopher Frog. Florida Gopher Frog. Southern Gopher Frog. Florida Frog-

RANGE

Check list. Range, Florida. Type locality Micanopy, Fla. — Stejneger
Barbour (1917, p. 36).

Rana capita LeConte. Range "Florida District." Cope. Type locality
Riceborough, Ga.

Supplementary records. Mr. Richard T. Deckert (1920, p. 26) reports
"an adult male, collected in May, 19 19, by Chas. E. Snyder, near Pinelands,
Hampton Co., South Carolina, and it is quite hkely that the species extends a
good deal further north and east than even this locality."

This record north of the type locality of Rana capita LeConte prompts the
query whether Rana capito is an isolated colony of the R. areolata group
separating R. aesopus into northern and southern colonies or whether the
more reasonable conclusion might not be that Rana capito and R. aesopus are
one species extending from Pinelands, S. C. far down into Florida. See dis-
cussion under Affinities.

Local Okefinokee records. In 192 1 we secured it at Chesser Island July
17, and on August 9 Mr. Francis Harper photographed one along the St.
Mary's River, Florida, six miles north of Macclenny (i mile east of Smith
Bridge). In 1922 we secured adults or tadpoles from Trader's Hill, Ga., for
several miles along the Trail Ridge to the big bend of St. Mary's River and
across St. Mary's at Hihiard, Fla., August 17, 1922.

Our records are along the Trail Ridge which extends north and south from
Georgia into Florida, the southernmost record of our trip being HilUard,
Fla., and northernmost between Chesser Island and Folkston, Ga. Residents,
however, report them from farther north at Braganza and Hickox, Ga.

GENERAL APPEARANCE

This species has had several apt characterizations. Cope remarks (1889,
p. 412) "This singular form may be known at once by the short and squat
form of the body as compared with the size of the head, resembling in this some
of the Australian Cystignathidae." Miss Dickerson (p. 194) writes "The
Gopher Frog of Florida is very different from other frogs in general appearance.
It is squat and toad-like, having an unusually large head, with a prolonged
muzzle, and eyes prominent and bulging even for a frog. Because of the
unusual length and breadth of head, this frog has a mouth relatively larger
than those of other North American frogs. These structural characteristics
give the frog so pecuhar an appearance that although it may agree with Rana
palustris and with Rana pipiens in colour and in being prominently spotted,
there would never be any difficulty in distinguishing it as a different species."

Plate XXXIV
Gopher frog {Rana aesopus)

1. Cypress pond, Hilliard, Fla. August 17, 1922.

2. Male croaking at base of bush, Coat -bet pond, Chesser Id., Ga. July 17, 1921

Flashlight.

3. Male, Chesser Id., Ga. July 2t„ 1921. Ventral aspect, x c.4.

4. Female, Chesser Id., Ga. July i, 1922. Ventral aspect, x 0.46.

5. Gopher frog at entrance of gopher turtle hole. Oak ridge, St. Mary's river, Fls

August 9, 192 1. Flashlight.

6. Eggs, stained, Hilliard, Fla. August 18, 1921. x 1.5.

• .••

Rana aesopus 335

MEASUREMENTS
(Recent Material)

Head to angle of mouth (28 mm.)— 1.32 (66 mm.) — 1.34 (68 mm. cf )
— 1.31 (82 mm. cf)— 1.58 (95 mm.cf)— 1.33 (95 mm. 9)— 1.2 (108 mm. 9 )
in width of head; head to rear of tympanum, .92 (28 mm.) — 1.07 (66 mm.) —
1. 13 (68 mm.c?") — 1.2 (82 mm.cf) — 1.32 (95 mm.cf) — 1.18(95 mm. 9 ) —
I. II (108 mm. 9 ) in width of head; head to angle of mouth ( )-3.o-2.6i-
2.5-3.06-3.06-3.0 in length of body; head to rear of tympanum 2.15-2.44-
2.2-2.34-2.55-2.36-2. in length of body; snout .5-.92-.93-.88-.88-1. 03-83
in first finger; snout .35-.84-.80-.63-.78-.61-.80 in fourth finger; snout .43-
.70-.57-.53-.60-.7-.50 in first toe; eye i. 74-1. 73-1. 4-1. 68-1. 54-1. 5-1. 7 in
snout; eye ( )-.93-.65-.83-.82-.7-.8 in tympanum; eye .875-1.6-1.3-1.67-
1.36-1. 5-1.4 in first finger; tympanym ( )-2. 7-3 -5-3 -3-2. 9-3 -3 7-3 -2 in
intertympanic width; tympanum ( )-i. 85-2. 1-2. 2-1. 9-2. 12 in snout;
internasal width .66-1.0-1.0-.94-.8-.94 in upper eyehd width; interorbital
width .64-1. 5-1. 16-1.0-1. 36-1. 0-1.25 in upper eyelid width; interorbital
width I. o-i. 62-1. 16-1.0-1. 45-1. 23-1. 33 in internasal width; interorbital
width ( )-4. 75-3. 8-3. 5-4. 5-4. 1-4. 5 in intertympanic width.

Forelimh: Forelimb 2. 33-2. 0-1.8-2. 05-2. 06-2. 1-2. 5 in length of body;
forelimb 2.75-2.75-2.9-2.8-2.82-3.0-3.4 in hindlimb; first finger i. 28-1. 08-
1.15-1.06-1.13-.94-1.21 in third finger; second finger i. 8-1. 3-1. 5-1. 6-1.7-
1. 37-1. 5 in third finger; second finger i. 4-1. 2-1. 3-1. 5-1. 5-1. 4-1. 25 in first
finger; third finger 1.1-1.2-.90-1. 06-1. 06-1. 15-.95 in second toe; fourth finger
1. 8-1. 09-1. 36-1. 45-1. 3-1. 58-1. 3 in third finger; fourth finger 1.2-.82-.72-
.82-.77-1.1-.64 in first toe; internasal width 1. 16-1. 84-1. 85-2. 14-1. 87-2. 19-
1.9 in first finger; internasal width .9-1. 5-1. 4-1. 4-1. 25-1. 56-1. 5 in second
finger; internasal width 1.5-2. 0-2. 14-2. 28-2. 12-2.06-2.25 in third finger;
internasal width .93-1. 7-1. 57-1. 57-1. 62-1. 3-1. 75 in fourth finger;

Hindlimh: length .84-.72-.63-.72-.73-.71-.75 in hind Hmb; tibia 2.54-
1. 9-1. 7-1. 9-2. 0-1.9-2. 1 in length; tibia 3.0-2.6-2.67-2.6-2.7-2.7-2.8 in hind
hmb; tibia 1.09-.94-.925-.93-.96-.9-.81 in forelimb; tibia 1.4-.9-.925-
.95-.91-.90- .90 in hind foot (without tarsus); first toe i. 66-1. 7-1. 75-1. 77-
1. 8-1. 65-1. 88 in second toe; first toe 2.-2. 5-2. 56-2. 55-2. 35-2. 52-3.1 in third
toe; first toe 3. 13-3.33-4. 18-4.3-4.0-3.4-4.5 in fourth toe; first toe 2.0-2.55-
2.25-2.44-2.4-2.17-2.87 in fifth toe; second toe i. 4-1. 45-1. 46-1. 3-1. 3-1.5-
1.64 in third toe; second toe i. 2-1. 45-1. 3-1. 3-1. 33-1. 31-1. 53 in fifth toe;
third toe i. 42-1. 33-1. 65-1. 5-1. 7-1. 4-1. 43 in fourth toe; third toe .82-1.02-
.87-.94-1.02-.86-.93 in fifth toe; fourth toe i. 55-1. 05-1. i-i. 05-1. 1-1.15-1. 14
in hind foot; fourth toe 1.1-1.16-1. 19-1. i-i. 2-1. 25-1. 25 in tibia; fourth toe
1.2-1.1-1.1-1.02-1.15-1.12-1.0 in forelimb; fifth toe i. 06-1. 3-1. 86-1. 7-1. 66-
1.6-1.5 in fourth toe; internasal width i.o-i. 4-1. 14-1. 28-1. 25-1. 4-1. 15 in first
toe; internasal width 1.66-2. 3-2. 0-2. 4-2. 25-2. 37-2. 12 in second toe; inter-
nasal width 2.33-3.4-2.9-3.3-2.7-3.6-3.5 in third toe; internasal width 3.33-
4.6-4.8-5.5-5.0-5.0-5.0 in fourth toe; internasal width 2.0-3.5-2.55-3.14-
3.0-3.12-3.25 in fifth toe.

336 Frogs of the Okefinokee Swamp

HABITAT

This species seems to be restricted almost solely to the burrows of the
Gopher Turtle {Gopher us polyphemus) which is common in the higher pine
barrens and sand hills along the swamp's eastern border. All the turtle burrows
might have frogs but these seem to be localized in colonies. Possibly, the
breeding habitats (cypress and other ponds) indicate the grouping of these
colonies or the animal associates of the gopher turtle may limit or terminate
whole or incipient groupings.

Occasionally the frogs live in other covers than turtle burrows. In one
colony near Chesser schoolhouse as we started to walk into the turtle area at
6:00 p.m., June 30, 1922, we heard something go into a small opening which
looked like a rat hole. The hole extended 18 inches into the ground and the
end was only 9-12 inches below the ground. Therein we found a female
gopher frog. Near another turtle's burrow we saw a similar hole with a
smooth worn spot about 8-12 inches away, the resting place of the frog. This
observation the residents have often made and two of them remarked about
this worn spot a foot or less from the hole.

On our previous trips 191 2-192 1, we have never found this species in the
Pocket nor on Billy's Island nor on any of the other islands within the swamp.
Like subterranean mammals (gophers, moles and shrews) subterranean
turtles (gopher turtles) and other subterranean amphibia (spadefoots) it
tends to avoid these situations which may be unduly flooded at times, or
where a shallow burrow may reach water level. On Chesser Island, where a
small narrow tongue of wooded swamp separates it from the mainland we
occasionally secure shrews, moles quite frequently, and spadefoots rarely.
Are gopher turtles and frogs also there? The gopher turtles occasionally
appear but are immediately sought by the inhabitants. 'Though the few
Gopher Turtles which reach Chesser's Island are soon killed off by the resi-
dents (not only because of damage to the peanut crop but also because their
burrows form too safe a retreat for hunted rabbits), the frogs remain, . . . ,"
(Harper). Whether the frogs come coincident with the gopher turtles, we
have not determined, but on July 17, 192 1, we made our only record of the
frogs on Chesser Island. About a dozen males were recorded croaking. There-
after neither in the summer of 192 1 nor 1922 were they found on this Island
where no gopher turtles are allowed, yet they are present as our one record
proves. The residents assure us that the frogs betake themselves to rat-like
holes, hollow stumps, holes beneath stumps, trees and logs. And I can well
believe that this species, were gopher turtles absent, might also resort to cray-
fish burrows (Gaige 19 14, p. 4), a retreat for R. areolata, a closely related
species.

Doubtless no other frog proves quite so typical of the higher pine barrens
unless it be the oak toad (Bufo quercicus). Occasionally the toad {Bufo
terrestris) the spadefoot (Scaphiopus holbrooki) and the swamp cricket frog
Pseudacris nigrita may be found in the same plant association. The other
animal associates are discussed under the enemies of the gopher frog.

Rana aesopus 337

"Gopher Frog (Rana aesopus). One male specimen was taken from burrow
No. 7 in the Cedar Creek locaHty on February 19, 1922. A Gopher Tortoise
(7 X 5 1/4 inches) was also taken from this burrow at the same time' Two
specimens were taken from burrow No. 10 in the Dinsmore locality on March
26, 1922. They were observed on two days just before sunset, resting in the
entrance about two feet down the burrow and they were not seen at other
times during the day. A Gopher Tortoise (11x8 inches) was taken from this
burrow at the same time. One specimen was observed about sunset at the
entrance of a burrow in the Brentwood locality on March 19, 1922. Two
specimens were taken from burrow No. 1 1 in the Dinsmore locality on April
12, 1922. They were observed several times resting in the burrow about two
feet down in the entrance just before sunset. A Gopher Tortoise (10x71/4
inches) was taken from this burrow, at the same time. In the Dinsmore
locality during the week of April 15, 1922, a pair of Gopher Frogs was ob-
served on several days about sunset resting just inside of the entrance of a
burrow. In the Cedar Creek locality on June 25, 1922, one specimen was
seen at 1 130 p.m. resting in the grass about 15 feet outside of the entrance of a
burrow and when disturbed made three hops and entered. The frog rested
motionless for a short interval of time between the jumps but the actions were
quick.

Gopher Frogs are not always found associated with Gopher Tortoises as
a Gopher Frog was taken in the Dinsmore locality on March 28, 1922 in about
6 inches of loose sand; another in about 8 inches of sand; and two specimens
were taken in a hole about two inches in diameter under a dead stump."
(Hallinan).

FIRST APPEARANCE

The earhest records we can find for the spring out coming are: March 18,
made by Prof. T. Van Hyning at Gainesville, Florida; and April 19 18 made by
N. R. Wood at Auburndale, Florida. Thomas Hallinan's records above are:
Feb. 19, Mar. 19, Mar. 26, Apr. 12, Apr. 15 and June 25, 1922.

GENERAL HABITS

This species is more common than usually held. In collections it is rare.
Most specimens have been taken at the critical period in the animal's life, its
breeding season when more frogs pay the toll than at any other time. In its
normal all-the-year-around habitat, the sand hills and turtle burrows they
are seldom seen unless one deliberately seeks them. They usually rest at the
mouth of the burrow, sometimes a foot or so down the decline or just within
the mouth. More rarely they may be one foot or more from the incline, or
be on little clear, smooth places 6 inches in diameter a short distance away from
a rat-like hole. One area we visited quite frequently. Usually from mid-
forenoon to mid-afternoon or later they were seldom out. For example, on
July 2, 1922, we visited this area at 3 :2o p.m. and found only one out but at
midnight we readily found eight. My journal reads: "Called again at 12
midnight. Francis's special friend not to be seen. Anna's friend was under

338 Frogs of the Okefinokee Siin)np

a log. The entrance a foot or more away because we had dug it back from
under the log. Suddenly something tumbled from under the log into the
hole before I realized what had happened. Francis found another at a hole
nearby. I found one and Miles saw three. One from i foot to i 1/2 feet from
the hole entrance proper. This one I caught. It is a female. Tried for one
or two more but did not succeed. They are not so strongly held by light as
are most Anura."

Nine days later, July 11, 1922, we have: "At 7:15 p.m. we started for
gopher frog area near Chesser School. Found 7 or 8 at entrance to holes or a
foot or two out. At one hole Miles saw a Peromyscus gossypinus. At another
I saw a Sigmodon. We captured three frogs. I caught one by hand. Could
have taken 3 or 4 more. Miles took the net without the handle. Interposes
it between frog and pole. Frog like a woodchuck makes for the hole and
jumps into the net. Sometimes this works, sometimes it does not. At first
I espied a frog at the entrance and tried to steal upon it from opposite the
incline. It heard me. Wonder if I could catch them with baited hook and
line. One was obliging and allowed us to photograph it. First one this year.
It turned around twice during the process then hopped in." With effort we
€Ould have photographed others. In 192 1, August 9, Mr. Harper stalked
another just north of Macclenny near the St. Mary's River on an oak ridge.
In both instances a log rests over the entrance. This is a favorite cover for
both frogs and turtles. The turtles seem at times to prefer to dig their hole
under fallen logs and the frogs often apparently prefer such gopher turtle
burrows or burrows with saw palmettos near by.

The gopher frog though often in colonies seldom in our experience lives
with another frog in the same burrow. This may be possible but not usual.
We never saw two in one burrow. R. F. Deckert (19 U, P- 3)> however,
writes that "Two or three frogs will inhabit the same turtle burrow," and more
recently (Feb. i, 1923) Thomas Hallinan (1923, pp. 19, 20) records several
instances of two in one burrow. His notes are definite, excellent and might
be given in toto.

Several authors from captive specimens have noted the color changes
of this form. Shufeldt (p. 1 54) holds "Gopher frogs are very clever at hiding,
and when doing so they change color at will, assuming a whitish tinge or brown
or deep purple, to intense black," while Miss Dickerson writes (p. 195) "The
general color of these frogs varies considerably. It may be grey or brown,
somewhat yellow or purple in tone. The spots may vary in size and in num-
ber. In any individual frog, the color changes are striking and rapid. The
frog may change from nearly black to white, through shades of brown or
purplish grey." Possibly educated frogs may thus operate with speed and
some of our captives showed different color phases and adaptation to different
environments, but I question whether a gopher frog might in its normal rela-
tions vary suddenly from white to black. The general average of frogs is of
the grayish hue. I never saw as sudden changes in this species as in Hyhdae.
A captured frog away from its burrow and placed on the neighboring sand,
may squat for protection and match its background beautifully. If disturbed

Rauo nesopus 339

it will start off with unexpected dash, travel a short distance and stop as
abruptly as its start was sudden. Several times I tried laying a female on
her back. Motionless she would lie. Then I would tickle the belly and she
would draw up her head and hind feet with the hind toes extended. Then I
would draw the head to within i to i 1/4 inches of the under side of the femur,
place the creature on its haunches. A ludicrous sight it was from the side or
front. After a minute or so of such posture, when we were in the midst of our
laughter, this inert idol would rapidly right itself and quickly scurry off with
short rapid hops. Indeed in the sandy saw palmetto oak ridges and pine
barrens coloration is its best asset if away from its burrow, its poisonous
secretions doubtless helpful and its quick dashes deceiving to the startled eye.

VOICE

This was heard positively on the evening of July 17, 192 1 and August 17,
1922, In the former instance about a dozen began croaking in a cypress pond
on Chesser Island before darkness came on. The fact that they did not begin
on the former day after a hard rain makes me believe these to be stragglers
in breeding, possibly tardy on this island. Later in the afternoon when
they began one of the residents called it "the monster." At first while
wading in the deeper water of the pond in the early evening we would hear
this mysterious note in amongst the cypress trees usually at their base or
on logs. The frog would espy us first and all we could determine was a
big splash doubtless made by a large frog. Finally we heard one in shallow
water in a tussock of sedge 2 feet across and i foot high and another near
the base of a cypress in deep water. By waiting for some time two of the
residents located them. They were truly R. aesopus. After two hours
work on Hyla gratiosa we sought out a group of gopher frogs we were hear-
ing in the shallower western portion of the pond. One was beside a cypress
tree in a depression of fibrous roots. After we had focussed on it it literally
crawled around to a different position near by. I put the light on its nose
and tried to push it back with my fingers. Then it leaped. At the base
of a pine on a pile of chips i 1/2 feet above the surface was another croaker.
After a time (i.e., after it had had light on its face a short period) it would
"bat" its eyes toward the buUseye. AVe secured two flashlight photographs of
it. Another male was in fibrous roots at the base of a black gum tree. A
fourth was at the base of a "Cassenya" (also "spice bush" pond spice (Mala-
poena geniculata) . Another was under a log where a tussock of sedge was over-
hanging. These frogs were not hard to catch when discovered but they were
shy. Often when first put under the light they would sink back in the fibrous
roots, depressions and covers. One (the croaker, PI. XXXIV, 2) was spread out
in the water among the spice bushes. The influence of nearby croakers seemed
to stimulate this one as it does others. We caught several, put them in a bag,
induced these to croak, and in this way our sprawled-out specimen responded
beautifully with his croaks. To induce the captives to croak we would swing
the bag round and round or rub them through the cloth or occasionally press
and hold in hand a male just ahead of the hind legs. We were sure we ought

34© Frogs of the Okefinokee Simmp

to find pairs and eggs in the pond the following day and evening. We dis-
covered neither but I am now sure the eggs found the following day were some
of the Rana aesopus eggs and not solely R. sphenocephala as I then pronounced
them. This fact alone showed how closely this species resembles the southern
meadow frog in this regard.

The preceding record (July 17, 192 1) was of frogs from an island in the
edge of the swamp where they do not normally live in gopher turtle burrows.
The second experience came August 17, 1922, on the Trail Ridge (oak ridge)
near Hilliard, Fla. After an excessive rainy period we were detouring on the
Dixie Highway when Mr. Harper heard spadefoots in some shallow pine barren
ponds. Later in the afternoon before and after 5 :oo p.m. I thought I heard
frogs on the higher open places in the pine forests. We suspected they were
gopher frogs and queried "Are they on the ground or at the gopher turtle
burrows or moving to their breeding places?" We never found one and one of
the party thought it a ventriloquial effect from a distant cypress pond. When
darkness came on this pond was suddenly resonant with gopher frog calls in
chorus. These chose similar perches to those described for 192 1. One was at
the base of a stump and above the water. Another was in the notch of a
floating log. A third at the base of a cypress. A fourth was beside a stump
and mostly sunk in the water. A fifth was at the edge of a small subsiding
pine. With all four of us searching we saw only one croak and all the rest
never responded when we were near. Several times Mr. Harper focussed on
males but we never secured the desired reaction. They were very shy. This
evening they frequented the edges of the pond where stumps, logs, and debris
were but were thickest in grassy and sedgy areas. These frogs called all night.
The chorus stopped at daybreak. Occasionally until 9:00 or 10:00 a.m. of
June 18 they would break out in a choral call, then subsided for a long period.

The vocal vesicles Shufeldt pronounces most unusual (191 7, p. 154), but
we doubt whether he or any writer who had captive specimens has ever seen
them at their maximum. This is possible at night in the field. In this regard
it is an exaggerated R. pi piens or R. sphenocephala with the vocal sac extending
almost to the groin. It constitutes the most striking development in vocal
sacs I have seen in North American Safientia.

"The croaking is a very remarkable performance. A lateral pouch is in-
flated on each side of the neck and keeps swelling posteriorly until it extends
halfway down the side of the body, attaining a size nearly equal to that of the
frog's head. The creature's form is then strongly suggestive of a bat-fish
{Cgcocephalus lespeitilio). The note — and likewise the inflation — lasts for a
second or so." It is a deep, hollow roll, somewhat intermediate between a
snore and a groan: yawhhhhh, yawhhhhh. At the start the intervals between
notes may range from ten seconds to nearly half a minute, but as the frog
warms to the performance, a croak may be given every two or three seconds.
The sound will probably carry a quarter of a mile." (F. Harper.) In 1922
both Mr. Harper and three more of us noticed the call might extend consider-
ably over a second's time.

Rana aesopus 341

When we first heard the note (July 17, 192 1) one of the residents thought
it a woodpecker but I recorded that it "seemed more hke R. sphenocephala
in tone" before I knew its close relationship to this form. At 7:30 a.m.,
August 18, 1922, several gopher frogs and southern meadow frogs were call-
ing. The call of the latter was 3 or 4 rasping croaks, then 3 or 4 clucks, i.e.,
dissimilar notes. The call of R. sphenocephala is like a rattle compared to the
uniform continuous roar of R. aesopus. To Mr. M. D. Pirnie it sounds like a
snore, Deckert also (191 4, No. 5, p. 3) states that "Its call is a loud deep
snore, ..." In chorus it sounded to Mrs. Wright and myself like the surf
or a deep gutteral roar (possibly a trill at times) like rolHng r's down deep in
the throat. We also likened it to a snore. The chorus seemed to go in waves
and have decided crests.

Deckert (1920, p. 26) records of a captive specimen that "When the interior
of the large glass jar in which it lives is sprinkled, and after this certain noises

are made, like rusthng paper, or water running from the tap, this frog "sings "

During the calls, which are repeated about every two seconds and are of from
three to five seconds' duration, the vocal vesicles over the arms are distended
into hemispheres about the size of large hazelnuts." We doubt whether these
gentlemen or any writer who had captive specimens have ever seen them at
their maximum.

MATING

Male {From life July 20, 1921). Sides of head, dorsum of limbs and sides
below costal fold French gray or lilac gray. Fore and hind legs with numerous
black bars. These bars broken at times or more or less speckled with French
gray. Costal fold from honey yellow to mustard yellow or buff (yellow to
greenish yellow depending on the degree of dark specking on it.) Another
similar stripe along upper jaw and over the arm insertion. All the tubercles
on the back with same color. Iris upper part black then buff yellow or honey
yellow over the pupil. Black in front and behind the pupil. Two lilac gray
areas beneath pupil separated by black. Between the costal folds about four
rows of black spots. Under parts white with hair brown or fuscous or pur-
plish black spots on throat and chin. Posterior faces of hind limbs purplish
vinaceous.

Female (From life, July 2, 1922). General color on the sides is French
gray, cinereous or pale purplish gray and some pallid purplish gray. These
colors become on the lateral fold, canthus rostralis, upper eyelid, beneath eye
and fold back of angle of mouth, cream buff or tilleul buff. Ground color of
back between lateral folds a combination of lateral fold color and side color.
Between the lateral folds are four more or less irregular rows of black roundish
spots. Below the lateral fold are 3 such rows to the rictal fold. Ground color
of body on dorsum of hind and fore limb with black cross bars. Undersides
of feet and hand and femur vinaceous lavender. Spots on the throat bister
to snuff brown. Those on sides of belly, army brown. Iris black with promi-
nent bar of colonial buff or tilleul buff above the pupil. Below the pupil may
be two small bars of body color, namely French gray, cinereous, pale purplish

342 Frogs of the Okefinokee Swamp

gray or pallid purplish gray. In another specimen two lower bars united.
Rest of eye black with a little dotting (irregular) of the two combinations
(upper and lower bars) of colors.

Structural differences. Some of the differences in males and females appear
summarized below:

1 . In describing the coloration we noted that "the males have yellows
on the folds, tubercles, iris, axilla and groin somewhat wherein the
females have little or no yellow. Possibly LeConte's description of
R. capita is from a male. This may need more rechecking.

2 . The male has the thumb enlarged and the female not, but the thumb
is not as enlarged as in some species.

3 . The males average smaller. The adult males externally are first re-
vealed at 68 mm. and reach to loi mm., while the females range from
77-181 mm.

4 . The antebrachium is very much wider in the male than the female and
this widening may even reach up to the brachium.

5 . Under Voice attention is called to the sacs which in Rana aesopus as
well as R. areolata may be immense and in alcohol may appear as ex-
tending far beyond shoulders. In life sometimes they look to reach
even farther backward.

6 . Some notes on alcoholic specimens are :

(a) 68 mm. cf. Thumb enlarged.

(b) 68 mm. cf. Thumb enlarged. Sac partially developed.
Very spotted on throat and pectoral region.

(c) 81 mm. cf . Thumb somewhat enlarged and slightly colored.
Sacs fairly developed.

(d) 85 mm. cf. Thumb enlarged. Sacs show.

(e) 87 mm. cf. Thumb somewhat enlarged and darkened.
Beautiful sacs half way to groin.

(f) 90 mm. cf . Thumb well enlarged, dark sacs well developed.
Amplexation. It is presumably axillary but we have recorded no pairs

of this species.

OVULATION

Habitat. We found it breeding in a cypress pond with no open center.
We found tadpoles in an open centered cypress pond, in an open centered
pond with bay tree circle, in an open pond with St. John's wort {Hypericum)
circle, in an open pond in pine woods or an old field, no special circle of vege-
tation, in an open pine forest pond at the edge of a wet savanna.

Period. We have long suspected that this form bred in the first half of
the year. In 192 1 we were within the swamp and missed this form until
July. In 1922 we arrived June 11. Deckert (191 4, No. 5, p. 3) writes: 'Tts
call can be heard from late February to the end of May. These frogs croak
only at night, and are then easily caught by the Hght of a lantern. The writer
has also collected this species during the breeding season in February near
Jacksonville, Fla., in 191 2 (Deckert, 1920, p. 26). It would seem then that it

Rana aesopus 343

bred from late February to August 17, the bulk of the ovulation comes before
summer or possibly it starts early and continues intermittently until Septem-
ber like its relative R. sphenocephala. The period of ovulation, as we ourselves
know it for eggs is from July 17-August 17. Mature tadpoles discovered
July i8-August 25 indicate that the eggs must have been laid in the late
spring at least, late April or in early May. We, therefore, derive a period
from late April to mid August.

Since the above evidence was considered, we found Prof. T. Van Hyning's
(1923, p. 68) pertinent note "On the night of March 18, while some of the
boys of the biological class of the University of Florida, Gainesville, Fla., were
collecting frogs, among other species taken, were . . . eight specimens of the
Gopher Frog, Rana aesopus (Cope) ....

"These and other species were all in a pond breeding near the University.
One of the party collected eggs, and brought living specimens which have
spawned since. . . ."

Temperature and humidity. The data for August 17, 1922, is as follows:
The maximum air temperature for the day before for nearby localities was
from 18-91°, average 87°, or for the day of the record 82-91°, average 86°, the
minima 69-72°, average 70°, and 69-74°, average 72° respectively. AtHilliard
on August 16, 1922, .56 inches of rain fell, on August 17, 3.47 inches or 4.03
inches in two days or 7.03 inches from August 16-21. For July 17, 192 1, we
derive 85-87°, average 86° air maxima for the day before and 83-88° for the day
of the record, average 85°, minima for the day preceding 69-7 2 °, average 70° and
68-74°, average 71°, the precipitation for day before was heavy as much as an
inch or more. At camp on July 17, 192 1, it was 75° at 7:30 a.m.

The tadpoles of the summer must have been from eggs laid the last of
May or the first of June, 1922. From May 25 to June 4 at Hilhard there
were 6 days of .90 iriches of rain or more, at Glen St. Mary's seven days, the
total at Hilliard was 13.33 inches, at Glen St. Mary's 11.93 inches. It would
seem these eggs must have been laid May 26-28 or May 30- June i when there
was as much as 4-6 inches of precipitation in three days. If these July and
August records be late breeders apparently an inch or more of rain is needed
to start the stragglers for breeding and air temperatures of 68-74°, average
71° or 81-91° maxima, average 86° seem to obtain.

Egg-laying process. This species we did not observe in the egg-laying
process. In its breeding habits it is so like the meadow frog we could readily
believe it lays its mass as does this species. Doubtless the oviposition when
once begun is completed in 5-15 or 20 minutes. Very evidently one position
is maintained throughout the process.

EGGS

Attachment, egg mass. On August 17, 1922, we found several masses of
gopher frogs eggs laid in a cypress pond. On the north edge in water 6-8
inches deep and 30 feet from the edge of the pond I found at the base of a
small cypress (10 feet high) and amongst some brush a mass attached to an
upright twig. It was one inch below the surface, was a phnth of eggs black

344 Frogs of the Okefinokee Swamp

above and cream below. The eggs looked larger and farther apart than are
those of Rana sphenocephala. Near by a large cypress stump in an open
grassy area where pine cones were on the bottom a large mass was attached
to a sedge stem. Its top was level with the surface of the water. The water
was 9 inches deep. The mass was 4x5 inches square and i 1/2 inches thick.
At first the mass impressed all of us as bluish and two of us independently
likened it to R. sylvatica in this respect. The lower pole may be cream at
first but it soon becomes white. The whole mass when turned over reveals the
same white mass impression R. sphenocephala and R. pi piens egg masses give.

On the southeastern side of the pond where Mr. Pirnie was watching the
frogs the previous night he also found two masses among young cypress and
attached to sedges was one mass 6x8 inches square and i 1/2-2 inches thick.
It seemed to be emarginate but when the lobes were drawn out the mass be-
came 12x4 inches square. The second mass was amongst "all spice bushes,"
(Malapoena geniculata) was also attached to sedges and was 4x5 inches
across. It was only i 1/2 inches below the surface.

On one occasion, August 21, in a pool filled with R. aesopus tadpoles we
found a suspicious mass of jelly 6x6x3 inches, an intact mass of jelly evelope
of a frog complement which I believed to be that of R. aesopus. On another
occasion July 28, 1922, in the grassy center of a very large open pond I found
an immense mass of eggs, then assumed to be R. sphenocephala eggs but they
were those of R. aesopus. The jelly envelope as in R. pipiens and R. spheno-
cephala tend to stay together some time after the egg has hatched.

The egg masses vary from 4x5x1 inches through 4x5x1 1/2, 6x6x2,
6x8x1 1/2 to 12x4x2 inches in dimensions. They may be attached to
grass, sedges, pickerel weed, or other aquatic plant stems, twigs and brush or
be free at times on the bottom as in R. pipiens or R. sphenocephala.

LeConte's comment on R. capita may have led workers astray from the
gopher frog because they did not know its breeding habits. Obviously (p.
425) his word that it "inhabits Georgia in the ditches of the rice fields" per-
tains to its short breeding habitat.

Egg description. In Rana aesopus the outer envelope ranges from 4.4 to
6.0 mm. in diameter, the usual range 4.8-5.4 mm., the average 5.3 mm., the
mode 5.2 mm.; the middle envelope is from 3.1-4.4 mm., the average is 3.9,
the mode 3.8; the vitellus is 1.8-2.4 mm., the average is 2.0 mm., the mode is
2.0 mm.; the vegetative pole is cream at first later white, the animal pole is
black. The rough measurements of some ovarian eggs of July 25 were 1.6-
1.8 mm. Possibly they were not fully developed. The vitellus and inner
envelope of R. aesopus is somewhat larger than in the meadow frogs (R.
pipiens and R. sphenocephala). On August 10, 1922, I divided the eggs of
the left side of a female in eight equal parts. One of these parts had 313 or
for the whole side 2504 or 5008 for the whole complement. The eggs look
larger than those of R. sphenocephala and farther apart in the mass.

Dangers. The dangers from drying up little affect the gopher frog in
contrast to some of the other Salientia of pine barrnes, e.g., the oak toad,
southern toad and pine woods tree frog. It chooses in general permanent open
ponds and cypress ponds.

Rana aesopus 345

HATCHING PERIOD

We have only one observation on the period of hatching. Eggs laid in the
night time between 8:00 p.m., August 16, and 9:00 a.m., August 17, 1922, in
a cypress pond hatched on August 21, 1922, or in about 4 to 4 1/2 days. On
the same date near by in an open shallow pond spadefoots began laying in the
day time where water was at least 80° F. or higher. These eggs hatched in
i-i 1/2 days. In mid-day the water was warm. We had no thermometers
but the water doubtless reached 90-95° or 100° F. The shade of the cypress
pond, however, lengthened the period for the gopher frog and the temperature
did not go above 70 or 80° F.

MATURE TADPOLE

Color description from life {August 1, 1922). At times the tadpole has a
very greenish cast. Color on top of the head oil yellow, olive green, olive lake
or old gold ahead of level of eyes. Back of eye level yellowish citrine or olive
citrine where there are no black spots.

About four irregular series of black spots between lateral line series of spots
of either side. Lateral line pores closely set and lateral line canal distinct to
the tip of the tail. A dark spot on each upper eyelid. A dark spot or spots in
interorbital space. Top of head ahead of eye with several fairly large spots.
Spots most distinct and numerous from nostril to base of muscular part of the
tail, being on the sides interspersed with light pinkish cinnamon or pinkish
buff. This color is present on muscular tail and slightly on upper and lower
crests as small clusters. This light pinkish cinnamon or pinkish buff merges
into the block color of the belly which is cream-buff, colonial buff, or naples
yellow. This block color breaks up in gill or bronchial region becoming clusters
of fine dots or spots. Below eye to either side of bronchial region is some
light vinaceous cinnamon or vinaceous fawn. Middle of bronchial region
without these colors but with a slight fine speckling of black lines.

Background color of the tail mignonette green. Upper crest and muscular
part with large spots of black. Lower crest without large distinct spots except
at the tip.

Iris rim around pupil orange or dandelion orange to light cadmium. Rest
of iris black with mignonette green or apple green fine dots.

General appearance. Tadpole large (84 mm.) full and deep-bodied.
Venter strongly pigmented so that viscera do not show through in life or in
preserved condition. Tail long, tip obtuse or acute. Dorsal crest in depth
sometimes equal to or greater than the depth of the musculature and extending
on to body somewhat ahead of the vertical of the buds of the hind limbs.
Spiracle sinistral, directed upward and backwards. Spiracle distinctly below
the lateral axis. Eye above or slightly on lateral axis, and near lateral outline
in dorsal aspect than mid dorsal line. Muciferous crypts distinct. The dorsal
row from either side of the insertion of the dorsal crest extends obliquely for-
ward and outward to join the principal lateral row which extends from above
the middle of the tail musculature to back of the eye. Here it forks, sending a

346 Frogs of the Okefinokee Sivanip

supraorbital branch far past the nostril to snout and an infraorbital branch
also to the snout and quite far below the nostril in its course. Below the
principal lateral series is a lower one on side around the spiracle but forward it
is not very distinct.

Mouth parts. Teeth 2/3. Edge of upper labium slightly larger than beak
and fringed with teeth. In either corner a row of teeth about 1/3-1/4 of the
upper fringe. The outer end of this lateral row usually (abnormal in figure in
this regard) does not extend beyond the end of the upper fringe. The median
space between lateral rows is one to two times either row, nearer the condition
in R. pipiens. The third lower row about equal to or slightly larger than
single row of lower labial papillae, 1/3-2/7 shorter than first or second rows
which are equal and larger than the beak. The tadpole is clearly of the H.
pipiens or R. sphenocephala type.

Measurements. Length of body (28-33.5 mm.) in tail (74-81 mm.) 2.15-
2.85, average 2.62. Width 13. 5-19 mm. of body in its own length 1.6-2.07,
average 1.8. Depth (12-15 mm.) of body .88-1.7 in body width, average 1.14.
Depth of body 1.5-2.4 in body length, average 2.01. Depth (10-18. 5 mm.)
of tail in length of tail 2.6-3.5, average 3.05. Muscular part (6-9 mm.)
1. 45-1. 9 in depth of tail, average 1.75. Spiracle 1.1-1.7 nearer base of hind
legs or vent region (14-16. 5 mm.) than the tip of the snout, average 1.36.
Spiracle 1.4-2.0 nearer eye than base of hind legs or vent, average 1.55. Eye
1. 0-1.3 nearer to spiracle (5.0-7.0 mm.) than tip of snout (6-9 mm.), average
1. 12. Nostril 1-1.5 nearer eye (2.5-5.0 mm.) than snout (3.0-6.0 mm.),
average 1.25. Mouth (3.0-5.0 mm.) usually .84-1.26 larger than internasal
space (2.8-5.0 mm.), average 1.06. Mouth contained i. 25-1. 66 (average 1.5)
in interorbital distance (4.5-8.0 mm.). Internasal space contained in inter-
orbital space 1. 3 -1. 9, average 1.57.

The dimensions of the largest tadpole are:

mm.

mm.

Total length

81

Spiracle to vent

14

Body length

29

Spiracle to eye

7

Body depth

14.0

Eye to snout

9

Body width

15-0

Eye to nostril

4

Tail length

530

Nostril to snout

6

Tail depth

iS-o

Mouth

5

Musculature of tail

8.0

Interorbital distance

8

Spiracle to snout

17.0

Internasal distance

4-

General remarks. For a long time there were two forms R. aesopus and
R. virgaiipes whose tadpoles we did not know positively. At that time there
were two tadpoles we were collecting we had not placed; one was not placed
until late in the season and proved to be the larva of R. heckscheri. The other
by elimination had to be R. aesopus or R. viigatipes. When these larvae began
to show four rows of spots on the back of the tadpole before hind legs were
well developed we were quite certain. But with the coming of the hind legs
and outpushing of the fore legs, the coloration of the dorsum of the adult

Rana aesopus 347

gopher frog was clearly foreshadowed. Although the mature R. aesopus
tadpoles are very much like those of R. sphenocephala they are too big for
the latter species. No doubt in their earlier stages they may cause workers
difficulties for some time to come.

LARVAL PERIOD

When we arrived on June 11, 1922, residents held that frogs of many
species were calling and active about two weeks before in very heavy rains of
that period. Inasmuch as this form is so strongly of the R. pipiens group in
breeding habits I believe it has about the same larval period, i.e., about 75 to
90 days. The transforming gopher frogs of August 25 to September 15 were
doubtless from eggs laid between May 25 to June 5. For example, at Hilliard,
Fla., straight east of Chesser Island and across St. Mary's River the precipita-
tion from May 25 to June 5 was 13.37 inchesor3.3oinchesMay 28,3. 2oonMay
31 and 2.80 inches on June i. In this same period Glen St. Mary south about
10 miles had 10.66 inches. This would mean a larval period of 85-^106 days.

TRANSFORMATION

Period. After the discovery of the suspected R. aesopus tadpoles, it was
a whole month before we were positive that these tadpoles of several open
ponds on Trail Ridge were actually R. aesopus larvae. For example our notes
would run somewhat as follows: "Aug. 9. Holt's pond, east of Starling Branch
Crossing. Found several of the tadpoles like those of Nigger Pond, Hypericum
fasciculatum pond and other Old Okefinokee Road ponds. They are very
large with no leg buds apparent or leg buds are very small. Tadpoles too big
for Rana sphenocephala J ^ On August 21, 1922, in one pond we secured them
at an advanced two-legged stage. Two days later, three-legged stages were
recorded. The illustration represents an individual with the right arm appear-
ing before the one of the spiracular side. On the same day one or two had
reached the four legged, long tail stage. On August 25 the short tailed four-
legged stage was reached. These were carried in a tin can on the running
board of the Ford on our northward journey and on August 27 and 28, 1922,
completed transformation. These are the only observations on actual trans-
formations. Many of the tadpoles of these open ponds probably would not
have transformed before September 15 or October i. It is very doubtful
that the tadpoles fromtheeggs of August 17, 1922, transformed before October
20 or November i. On October 26, 1922, Thomas HalHnan found a specimen
just past transformation. Transformation may then be from August 15-
November i. But if this species begins breeding in late February or March
Deckert's transformation must come by May i or before. What puzzled
us was our absence of any records of transformed individuals from June 1 1-
August 25. We left August 25 and, therefore, had no opportunity to watch
the dispersal of the newly transformed frogs, their choice of habitat, etc.

Size. The transforming frogs of August 22-27, 1922, from a pond near
Chesser School (see U. S. Geo. Survey quadrangle), Ga., measured 28, 32.5
and 33 mm. respectively. On the basis of tadpoles approaching transforma-

348 Frogs of the Okefinokee Sivamp

tion we have one note on transformation as follows: "28-35-(?) mm. In the
American Museum material (Nos. 5930-32, 5896, 5897) are five specimens
from N. P. Fry of Eureka, Florida. These are 32, 27, 28, 33, 36 mm. respec-
tively and from near Arlington, Florida, Thomas Hallinan takes a specimen
31 mm. These all must be at or near transformation. Possibly transforma-
tion extends from 27-35 or 36 mm. It makes its transformation size com-
parable to the upper part of the transformation range of Ra7ia pipiens (19-28
mm.) or of Ra7ia sphenocephala (19-33 mm.). It is evident if we had enough
material that the mode and average of Rana aesopus would be higher than
that of Rana pipiens (22 mm. and 24 mm.) or of Ra7ia sphenocephala (25 mm.
and 23 mm.).

GROWTH

We have only transformation and adult sizes. The intermediate stages we
missed. From analogy one would believe that at least 3 years were required
for their maturity and possibly 6 years for the maximum sizes. Deckert
(1914, p! 3) writes ''The one-year-old frogs of this species I have repeatedly
caught, wandering from the high ground and its burrows into the swampy
meadows below, where the insect hfe is more abundant. On being discovered,
they will not dash away with long leaps, like other frogs, but squat close to
the ground, motionless, like some toads, and are then easily picked up."

This species apparently transforms at 28-33.5 mm. Our adult material
begins with sizes of 66, 68 mm., 73, 82, 85, 95, 95, 108 mm., and it seems to
sort into groups of 68-73 mm., 82, 95 mm. and 108 mm.

In Levy Co., 1892, J. Hurter (U. S. N. Mus. Nos. 57533-35576-78)
collected 62, 69 mm., 71 and 74 mm. specimens — apparently one group 62-74
mm. In July, 1893 (U. S. N. Mus. No. 20513) H. G. Hubbard collected a 65
mm. specimen and on June 20, 1894, two specimens (U. S. N. Mus. Nos.
21702, 21703) 71, 73 mm. respectively or one group 65-73 mm. At Auburn-
dale, Fla., N. R. Wood secured in 1917 (No. 59413) a 74 mm. specimen and
another in April, 19 18, 49 mm. in length, apparently two groups. The type
is 52 mm. and they have a specimen 83 mm. Boulenger's material was as
follows: 68, 73, 77, 80, 82, 84, 85, 88, 91, 93, 95 mm. respectively. One lot taken
at same place is 68, 84, 85 and 93 mm., two groups 68 and 84-93 mm. ; another
lot is 73 and 95 mm., two groups; and another lot 80 mm., apparently one group.

The American Museum material seems to sort: 27-36 mm.; 49-55 mm.;
62-76 mm.; 78-85 mm.; 90-102 mm.

In the Museum of Comparative Zoology series were 12 specimens taken
in 1919 at Orlando by Dr. Thomas Barbour. These are 58, 61, 63, 64, 67, 67,
68, 70, 70, 71, 76, 93 mm. The same year Dr. Barbour secured several 10
miles north of West Palm Beach, Florida. They are 82, 86, 89, 89.5, 90, 94.
Other records they have are:

No. 3574 Fruitland Park, A.G.Reynolds, 1914, 40 mm.

5137 Gulfport, Fla. " " " 1919, 52 mm.

7081 " " " " " 1920, 41 mm.

5938 Sebastian, Fla., George Nelson, 1919, 74 mm.

4907-10 New Smyrna, Fla., March 1919, 79, 86, 89, 91 mm.

Rana a e so pus 349

This material might be interpreted as 40-52 ist-year-olds; 58-64 2nd-year-
olds; 67-76 3rd-year-olds; 79-86 mm. 4th-year-olds; 89-94 sth-year-olds.

Our material apparently fell into groups as follows: 28-33 ™ni- ^t trans-
formation and adult modal groups of 68, 85, 95 and 108 mm. Does it mean
four or five different years? The whole material of the U.S. National Museum,
American Museum, Boulenger's eleven specimens and our own might be in-
terpreted as falling in groups 28-38-(?) mm. at transformation; 38(?)-52 mm.
first-year-olds; 52-65 mm. for 2-year-olds; 66-77 mm. 3-year-olds; 78-88 4-
year-olds; 89-102 mm. 5-year-olds; 102-108 mm., 6-year-olds.

FOOD

The writer has made few direct observations on the feeding habits.
Much which has been written has been on captive specimens. Shufeldt ('17,
p. 155) asserts that "Gopher frogs feed upon small birds and insects, but most
commonly upon toads, the last giving them no end of discomfort on account
of the acrid secretions from their parotid glands. Indeed after a meal upon
a big toad, this frog has been observed to go through a series of somewhat
extraordinary spasmodic movements, frantically trying to clear its mouth
with its fore feet of the semi-poisonous juice the batrachian he had endeavored
to swallow had squirted into it. So big is the mouth of this gopher frog that
it can manage to swallow a full-grown specimen of our common toad — a feat,
however, that it in no way appears to enjoy."

Miss Dickerson holds (p. 196) this frog seems especially fond of toads as
an article of diet. It ejects the poison from the mouth as soon as it has swal-
lowed the toad. A toad of surprisingly large size can be managed, owing to
the unusual development of the jaws and throat of this species. If the toad is
too large to be swallowed at once, so that the poison can be sent out through
the mouth almost immediately, the frog gives up the attempt and disgorges
the toad after about fifteen minutes. This time, however, is sufficient to
allow some action of the poison to take place, and the frog shows many signs
of discomfort. It has convulsive movements of the muscles, it leaps blindly
upward, and finally, lowering the head and opening the cavernous mouth,
uses the hands in frantic efforts to remove the irritation from there. During
several weeks of captivity, Rana aesopus fed almost wholly upon toads, of the
species lentiginosus, woodhousei and fowleri."

Deckert (1920, p. 26) records that his "specimen is in fine condition and
not at all shy, taking meal worms, roaches, and spiders and occasional earth-
worms from one's fingers, if not approached too suddenly."

In 1920, Dr. Thomas Barbour (1920, p. 55) presented before the American
Society of Ichthyologists and Herpetologists notes on this species. His
observations on the "dispersal and the habits of feeding on the small oak toad,
Bufo quercicus," I did not hear and they are not expanded in the abstract of
the proceedings of this meeting.

ENEMIES

The gopher turtle's burrow seems to be the permanent or temporary resort
of most of the animals of the drier pine barrens. Many smaller forms like the

350 Frog a of the Okefinokee Swamp

oak toad (Bufo quercicus), the six-lined lizard (Cneniidophorus sexlineatus)
glass snake (Ophisaurus ventralis), smaller snakes, peculiar insects, cotton
mice (Peromyscus gossypinus gossypinus) enter in and out of the burrows, but
it is not especially conceivable that they would seriously injure large adult
gopher frogs. If the young frogs live also in the burrows they might succumb
to some of the forms above. Those who like to think of the pine snake, the
gopher snake, the diamondback rattler, the cottontail rabbit, the gray fox, the
skunk, the burrowing owl all living in amity together will no doubt point out
that the snakes and possibly some of the others feed on warm-blooded prey.
Experience proves, however, that the gopher snake may eat glass snakes as
well as rats, that the pine snake and diamondback may take cold-bloods
occasionally and that the carnivorous mammals would not wholly avoid frogs.
The shell of the turtle makes it almost immune from attack. The gopher frog
has two defences, one its poisonous secretion and the second its ability to
crowd down in the sand of the burrow. The first defence we can hardly be-
lieve wholly effective with all snakes and possibly some carnivorous mammals,
the second defence is not as strong proportionally in the frog as in the turtle.
One wonders if the turtle be in any way a protection or shelter to the frog?
One male, 95 mm. in length has no left hind leg except for the thigh. What
enemy took the rest?

In its breeding habitat if it be in an open pond the water snakes prove
serious foes at night and in cypress ponds both water snakes and moccasins
are searching for all frogs.

This species has as distinctive an odor as Rana palustris or Ra7ia septen-
trionalis or R. virgatipes. The containers in which we have kept gopher frogs
may be frothy as we have experienced with R. palustris. We have never tried
putting other species of frogs with them to see how the other species would
withstand these secretions.

AUTUMNAL DISAPPEARANCE

Our own personal acquaintance with it is from July 17-August 9, 192 1,
June 14-September I, 1922, when our party encountered it in the Okefinokee
region. The type of Cope's Rana areolata aesopus U. S. National Museum
(No. 4743) was taken by Dr. Bean Dec. 23, 1886. In the American Museum
material is one small specimen (31 mm.) taken "in the grass about 40 yards
from a cypress swamp near Arlington, Florida" by Thomas Hallinan on
October 28, 1922. This is in the very year of our experience with the same
species.

AFFINITIES

I believe first that Rana capito LeConte is Rana aesopus Cope and secondly
that Rana aesopus thus understood {R. capito and R. aesopus) is closely
related to R. spheiiocephala and R. pipiens.

First, I have long suspected that R. aesopus was R. capito. Two summers'
acquaintance with the so-called R. aesopus in a region intermediate between
Riceborough, the type locality of R. capito and Florida (range of R. aesopus)
confirms this belief. "A few days ago I tried an experiment on my three as-

Plate XXXV
Gopher frog (Rana aesopus)

1. Gopher turtle hole, Trail Ridge, Folkston, Ga. July 5, 192 1.

2. Male at base of pine, Coat-bet pond, Chesser Id., Ga. July 17, 1921. P'lashlight.

3. Male on the sand, Chesser Id., Ga. July 23, 192 1.

4-6. Series of tadpoles, Chesser Id., Ga. Aug. 22, 23, 1922. x 1.2.

Rana aesopus 351

sociates of the summer of 1922 (Mrs. A. H. Wright, Messrs. F. Harper and
Miles D. Pirnie). I showed them LeConte's figure of Rana capita with the
label hidden and gave no intimation of locality at all. One instantly pro-
claimed it the gopher frog (Rana aesopus) on general appearance. Another
called it the gopher frog because of its waist and general appearance. The
third thought the figure a gopher frog but felt it too slim in the waist until
when shown a photograph of a gopher frog (Ventral view July i, 1922,
PI. XXXIV, Fig. 4) in a similar position she pronounces the two as one without
reservation." All four of us who have called upon gopher frog adults at all
times of day and night feel the two too close to be considered separate forms.

To be sure some may yet prefer to make R. areolata capita the northern
subspecies from Riceborough, Ga., northward and may restrict R. areolata
aesopus to the southern form. I see no real difference in the forms nor use for
this interpretation. Rana capita was described in 1855 by LeConte (1855,
p. 425) from the material secured in the ditches of the rice fields of Riceborough,
His description reads hke the Gopher frog {R. aesopus). In 1886 (1886, pp.
517, 518) Cope gave the four forms of R. areolata as R. a. areolata Baird &
Girard (Austroriparian region R. a. circulosa Rice & Davis (North Central
Eastern region, Illinois), R. a. capita LeConte (Floridan district) and a new
form R. a. aesopus Cope (Florida). Observe that Cope speaks of R. a. capita
as of the Floridan district. The combined ranges of R. capita and R. aesopus,
therefore, extend from Florida to South Carolina in the lower coastal region,
and we thus see R. capito more in agreement in range with other Floridan forms.

Cope (1889, p. 409-416) uses the same groupings of four subspecies and
and writes "This well-marked species (R. areolata) is related to the R. palus-
tris, but is easily distinguished."

We strongly incline toward the group which hold Rana aesopus separate
from Rana areolata and equally believe Rana capito and Rana aesopus the
same, but until more material is collected in Georgia and Carolinas, we little
wish hurriedly to make any changes from Rana aesopus to Rana capita,
though some may hold more or less rightly that consistency dictates such
action on our part.

Our second consideration that R. aesopus is closely related to R. pipiens
and R. sphenocephala does not militate against some relationship with R.
palustris but R. palustris is not the nearest relative. We have silhouettes and
other photographs of the gopher frog that remind one very much of R. pipiens,
and R. sphenocephala or even R. palustris. Like Rana pipiens and R. spheno-
cephala and unlike Rana palustris, the gopher frog has a plinthlike egg mass,
black and white eggs, similar breeding place (with R. sphenocephala), tadpoles
very similar and lateral vocal sacs. R. aesopus has vocal sacs extending to the
groin of either side, R. sphenocephala has them as spheres and R. palustris is
without them. R. aesopus and R. sphenocephala egg masses and tadpoles are
quite similar and perplexed us for a long period.

In breeding habits it did not need to be strongly differentiated from R.
pipiens and R. sphenocephala as these resort to the same type of place for
ovulation.

352 Frogs of the Okefinokee Swamp

In its non-breeding behavior, (protection, food, etc.) it appears most
peculiar. It had to adapt itself to the arid pine barrens and sandy oak ridges
wherein the only common moist retreat for a non-burrower is the gopher
turtle hole.

BIBLIOGRAPHY
1920 Barbour, T. Herpetological Notes from Florida. Copeia July 31, 1920, No. 84.

PP- 55-57-

1886 Cope, Edward Drinker. Synonymic List of the North American Species of Bufo

and Rana, with descriptions of some new species of Batrachia, from specimens in

the National Museum. Am. Phil. Soc. Proc, Vol. 23, Dec. 1886, No. 124, pp. 514-526.

1889 . The Batrachia of North America. Bulletin U. S. Nat. Mus. No. 34, 1889,

PP- 1-525-
D ■

1914 Deckert, Richard F. Further Notes on the Salientia of Jacksonville, Fla. Copeia
April 15, 1914, No. 5, p. 3.

1920 . Note on the Florida Gopher Frog. Copeia March 25, 1920, No. 80, p. 26.

1906 Dickerson, M. C. The Frog Book, pp. 193-196.

1914 Gaige, Helen Thompson. A List of the Amphibians and Reptiles Observed in Rich-
land County, Illinois, in May 1913. Copeia October 15, 1914, No. 11, p. 4.

1923 Hallinan, Thomas. Observations Made in Duval County, Northern Florida, On the
Gopher Tortoise (Gopherus polyphemus). Copeia February i, 1923, No. 115,
p. 11-20.

1855 LeConte, John. Descriptive Catalogue of the Ranina of the United States. Dec,
18, 1855. Proc. Acad. Nat. Sci. Phila., Vol. VII, 1854, 1855, Phila. (1856), pp

423-431-
1917 Shufeldt, R. W. The Florida Gopher Frog. Aquatic Life. August 19 17, Vol. 11,

No. 12, pp. 153-155-
191 7 Stejneger, Leonhard and Barbour, Thomas. A Check List of North American Am

phibia and Reptiles. Cambridge 191 7, pp. 1-125.
1923 Van Hyning, T. A Collecting Note on Florida Batrachians. Copeia May 20, 1923

No. 118, p. 68.

Rana clamitans Latreille

(PI. Ill, Fig. 2; V. Fig. 11; VIII, Fig. i; XI, Figs. 3, 4; XIV, Fig. 7; XVI, Figs. 4, 5; XVII;
XXXVI; Text Figs, i, i6)

COMMON NAMES

Green Frog. Pond Frog. Spring Frog. Bullfrog. Bawling Frog. Yellow-
throated Green Frog. Belly Bumper. Bully. Screaming Frog. Black Frog.

RANGE

Check list. Range: Eastern North America. Canada to Florida and
Louisiana, west to Michigan, Illinois and Arkansas (Stejneger & Barbour

1923, P- 35)-

Supplementary records. (Not sought).

Local Okefinokee records. We have taken this species on Billy's Island,
Chesser Island, Mixon Hammock, and in ponds along St. Mary's River and
in ponds beside the Folkston-Moniac road and in edge of cypress bay along
the eastern edge of the swamp.

GENERAL APPEARANCE

To the general lay it is a "yellowthroated bullfrog" and is usually confused
with its bigger congener, the bullfrog. In Copeia (1916, pp. 53, 54) R. F.
Deckert and Fowler 19 18 (Copeia, p. 84) reported a green frog which saw an

Ratia clamitans 353

albino. The first writer noted the scarcity of albino frogs. I have only seen
in the field one albino green frog, one albino meadow frog and Bufo americanus.
I have received two specimens of albinistic tadpoles of Rana heckscheri.

COLORATION OF SPIRIT SPECIMENS (1912)

(Spirit specimens of 191 2). On the back the color may be from deep
grayish olive through chaetura drab or mummy brown to olivaceous black or
almost black; in the young the colors incline the lighter side of the above
range and in these there are two, at times three prominent cross bars on the
femur and tibia, also cross barring on the tarsus and outer surface of the foot
and indistinctly on the forelimbs. In many of the adults these bars or spots
do not show at all or are faintly indicated; in a few they appear but may be
broken up. In the adults the small dark circular spots of the young are
absent or seldom present and the upper parts are almost a monochrome. In
most of the adults a black oblique bar extends from the angle of the mouth on
to the brachium and the back border of the forelimb has a prominent black
bar on the brachium and antebrachium; in all the adults the upper jaw ahead
of the eye is Hke the general dorsal color except for the snout which at times
is spotted with white spots; the whole edge of the lower jaw is a series of con-
spicuous alternate blotches of yellowish white or white and brown or black.
Except in very black individuals there is a yellowish white or white line from
beneath the tympanum (rarely from beneath the eye) over the arm insertion.
Tympanum brownish with a central disc of body color; the posterior faces of
the thigh are mottled with black and the anterior faces of the thigh less so;
the inner edge of the tibia has three or four short cross bars; also along the
outer face of the tarsus is a broad black band with a yellowish white line at
its lower border. This black band usually runs on to the two outer metatarsals.
In all the specimens are the more or less characteristic dark blotchings of the
sides, these spots being partly on the dorsal color and partly on the light
ventral color. The ventral color is white or yellowish white but is not im-
maculate as in our northern specimens. In many the under parts are heavily
marbled or reticulated with black, being the strongest on the throat and
upper breast regions and on the under sides of the thighs and legs. In some
where quite light beneath the reticulations or markings are faintly outlined
over the whole venter. In one specimen, a female, the throat and upper
breast region is almost solid black. In our northern forms when transforming
or just transformed, the frogs are very spotted or dotted on the throat and
under parts of the legs, but in the adults these marks are more or less lost.
In these southern forms the young are even more spotted on the venter and
the process consists in an intensification of spotting the reverse of the north-
ern tendency.

STRUCTURAL CHARACTERS (WRITTEN IN 1 91 2)
In none of these forms is the skin rough to the touch. On the dorsal sur-
face there are fine round mamillations, most coarse on the sides; the head is
without these and is merely pitted, the most rugose part being the outer half

354 Frogs of the Okefinokee Swamp

of the upper eyelid. On the ventral parts the skin is smooth except on the
buttocks which are decidedly pustulate and on the throat which is sometimes
lightly mamillate. Top of the head smooth, plane or rounded. A broad
short fold of skin on which is a yellowish white stripe extends from the angle
of the mouth over the arm's insertion. It is usually broken just ahead of the
arm by the fold which comes down the caudal border of the tympanum and
circles in front of the arm's insertion on to the breast. Sometimes the ends of
these last folds are connected across the breast by a faint fold or indentation.
The main fold with which this post-tympanic ridge is connected usually runs
from the eye backwards two-thirds of the distance to the hind limbs. At its
end it may break up or sometimes reach only halfway to the hind limbs. In
two small males these costal folds run little back of the tympanum and soon
cease. First and fourth fingers about equal. Third finger longest, subarticular
tubercles moderate or small; metatarsal tubercle small, third and fifth toes
about equal; last two phalanges of the fourth toe free, not webbed; tongue in
some broad and in others quite narrow, forked behind in all the specimens,
free for its posterior half and along edges of the anterior half; vomerine teeth
between inner nares or slightly behind line of nares, small, usually obliquely
set and closer together.

MEASUREMENTS
(Recent Material)

Head to angle of mouth i.i (28 mm.)-i.24 (56 mm. d^)-i.2 (82 mm.
cf)-i.2 (95 mm. d^) in width of head; head to rear of tympanum .83-
.92-967-1. 01 in width of head; head to angle of mouth 3.1-2.82-3.28-3.05 in
length (snout to vent); head to rear of tympanum 2.33-2.15-2.64-2.53 in
length; snout .7-.9-.916-1.0 in fourth finger; snout .7-.8-.83-.73 in first toe;
eye i.i-i.25-i.5-i.36in snout;eye .625-1.12-1. 62-1. 27 in tympanum; eye .77-
1.12-1.5-1.36 in first finger; tympanum 2. 4-1. 33-1. 3-1. 25 in inter-tympanic
width, tympanum 1.4-1.11-.92-1.07 in snout; internasal width 1.0-.91-1.0-
1.14 in upper eyelid width; interorbital width .875-1.67-1.40-1.6 in upper
eyelid width; .875-1.71-1. 4-1. 4 in internasal width; 2.1-3.4-3.4-3.5 in inter-
tympanic width.

Forelimb i. 86-1. 8-2. 0-1.75 in length; 2.72-2.77-2.7-2.71 in hind limb;
first finger i. 2-1. 33-1. 33-1. 26 in third finger; second finger 1.57-1.5-1.45-1.35
in third finger; i. 28-1. o-i. 09-1. 07 in first finger; third finger i.o-i.o-i.i 2-1.0
in second toe; fourth finger 1.0-1.14-.91-1.0 in first toe; i. 56-1. 7-1. 45-1. 26 in
third finger; internasal width i. 28-1. 5-1. 71-2. 14 in first finger; 1.0-1.25-1.57-
2.0 in second finger; 1.58-2.0-2.3-2.7 in third finger; 1.0-1.16-1.57-2.14 in
fourth finger.

Hindlimb: length i. 46-1. 53-1. 33-1. 53 in hind limb; tibia 2.0-1.86-2.0-1.9
in length; 2.92-2.77-2.7-2.92 in hind limb; i. 07-1. o-i. 0-1.08 in forelimb;
1.07-1.64-.902-.96 in hind foot; first toe i. 56-1. 5-1. 8-1. 72 in second toe;
2.28-2.25-2.2-2.54 in third toe; 3.14-3.75-3.5-3.91 in fourth toe; 2.0-2.56-
2,7-2.72 in fifth toe; second toe i. 45-1. 5-1. 22-1. 42 in third toe; 2.0-2.5-1.94-
2.26 in fourth toe; i. 27-1. 70-1. 22-1. 52 in fifth toe; third toe i. 375-1. 66-1. 59-
1.59 in fourth toe; .875-1.14-1. 22-1. 07 in fifth toe; fourth toe 1.36-1.1-1.05-

Rana clamUans 355

I.I I in hind foot; i. 27-1. 03-1. 17-1. 16 in tibia; i. 36-1. 03-1. 17-1. 25 in fore-
limb; fifth toe 1. 57-1. 46-1. 3-1. 48 in fourth toe; internasal width i. 0-1.33-
1. 42-1. 57 in first toe; 1.57-2.0-2.57-2.7 in second toe; 2.28-3.0-3.14-3.85 in
third toe; 3. 14-5.0-5.0-6. 14 in fourth toe; 2.0-3.4-3.85-4.14 in fifth toe.

HABITAT

Suitable ponds whether on the higher banks of St. Mary's River under
oaks or in overflow and cutoff pools with river swamp flora (Thompson's
Landing) , or whether they be the fringe of cypress and gums of a cypress pond
(Long Pond) or roadside ponds, sphagneous (Spanish Creek Woods) or not or
ponds in hammocks (Billy's Island) — all are used. The factors of pond and
shade both seem to enter. They were most abundant in chorus in the island
edges of the cypress bays or their counterparts, the fringe of trees around a
cypress pond. We also found them in the "dreens" (Cypress Bay) between
Chesser Island and the mainland.

In our experience with this species it is neither common within the swamp
nor outside the swamp. Probably it is easier of observation in ponds outside
the swamp or on the swamp's border but we are not prepared therefore to
say it is more common outside the swamp.

FIRST APPEARANCE

The earliest record we have is April 28, 192 1, but it must appear earlier.

GENERAL HABITS

In 191 2 we made the following observations regarding their general habits:
"Its habits in the swamp quite generally agreed with its known habits else-
where. It was found both outside and inside the swamp. All our captures
were from ponds and its tadpoles were found in the more permanent ponds or
swamp's cypress thickets on the islands. We seldom saw it on the prairies or
along the cypress runs where the common form was Rana grylio."

VOICE

The voice of the green frog is quite generally known to naturaHsts. Miss
Dickerson (1906, p. 199) says "it is explosive, prolonged, and low-pitched and
is likely to be repeated five or six times in succession. When we hear it given
with less than its usual force, we may be deceived into thinking that we hear
the drumming of a woodpecker. It may be imitated very well by cutting on
a table, with heavy shears, some kind of coarse resisting cloth. Or it may be
imitated less well by tearing heavy cloth in a jerky fashion. It resembles the
croaking of the Pickerel Frog, but has more strongly accentuated notes in it."

"The explosive character of the sound will be better understood from
watching the frog when he is croaking. He works hard; there is no appearance
of external vocal pouches (as in the Leopard Frog, Fig. 9), but the yellow
throat and the sides expand with such force as to jerk the whole body forward.

356

Frogs of the Okefinokee Sivamp

Slowly the distended parts sink in as the vigorous sounds proceed. Then the
throat and sides swell out again, and there follows another explosion of sound.
And so on over and over. ..."

Overton (1914, p. 37) writes of the green frog call as follows: "Its song is
an explosive note resembling that made by plucking the string of a bass violin.
It swells out its throat and cheeks during its song."

A resume of the croaking for 192 1 and 1922 follows:

1921
Entered swamp.
One male croaking.
Heard one or two lone males.
An occasional male calling.
One or two calling at 9:30 p. m.
Tried to photograph a few calling males.
One or two calling tonight. Air 70°
Green frogs croaking.
Quite a few calling 10:00 p. m. Air 77°.
Several calling.
An occasional R. clamitans.
Hear green frogs.

Hear no end of green frogs. Hear some in the forenoon.
Green frogs croaking their greatest.
Mainly green frogs calling in some ponds.
Several green frogs calling.
Many calling and several egg masses.
Great chorus last night.

Green frogs are now croaking in real earnest.

In east side of swamp no end of Rajia clamitans in early forenoon.
Hear green frogs. Eggs found.
Last record we made for calling.

1922
We encamped.
First green frog heard.
Hear a few in cypress pond.
Hear several.
One heard in cypress bay.
At 6:00 p. m. several calling.
One or two heard 7:00-9:00 a. m.
In early forenoon commonly heard in cypress bay.
One heard near camp. One heard about 2 :oo p. m.
A few in bay.

Heard a few in center of swamp.

Heard a few on St. Mary's River at Camp Pinckney.
Hear several.
Still calling on prairie.
One calling in mid afternoon.

When temperatures range in maxima 8o°-88° and minima S9°-^7° they
begin sparingly but when maxima reach 90° or more and minima 68°-75° they
begin to reach chorus stage, e.g., in 192 1 the choruses were greatest from July
2-1 1 when the temperatures were: maxima 88^-94°, average over 90° and
minima 69°-74'', average 70°.

April

23-

"

28.

May

14.

"

16.

"

18.

"

19.

"

21.

• "

22.

"

24.

"

26.

June

3-

"

8.

"

25-

July

2.

3-

"

4-

"

5-

"

II.

"

12.

"

13-

"

IS-

Aug.

19.

June

12.

"

22.

"

25-

"

27.

July

I.

"

4-

"

7-

"

10.

"

27.

'\

28.

Aug.

3&4

"

II.

"

15-

"

31-

Sept.

5-

Rana damitans

357

In 1922 the day we most commonly heard them was July 10 when some
of the maxima was 88°-93*' and minima 68°-7i° for nearby stations. On
August 31 they were calling on the prairies, temperatures maxima 88°-9i°,
minima 63 "-64°. The last date we have is September 5 but they doubtless
call much later in the season.

The late June or early July choruses came mainly at night, but often ex-
tended far into or throughout the day as well. Occasionally one will hear them
at mid afternoon, mid day and mid afternoon before chorus stage is reached.
Our record for September 5, 1922, is a mid afternoon call. F. Harper char-
acterizes the call as follows: ". . . its twanging bass note is heard at consider-
able and probably irregular intervals, of, say half a minute or more. Ordinarily
a single clung or cHung is given; but sometimes there is a rather rapid series
of several notes: chmg-clung-clung-dung or c'tung.'' The note is not loud
nor has it far reaching carrying effect. The male rests on a mat of vegetation
at the surface or amongst grass, or aquatic plants or freely sprawled out on
the water. Usually the hind quarters are slightly submerged. It is extremely
difficult to secure a good flashlight of its throat when slightly inflated. In the
first place it does not swell out its throat as far as in Rana grylio, and its call
is usually one note of very short duration. Furthermore when under observa-
tion long intervals often elapse between croaks. All in all it is one of the more
difficult frogs to catch at the full inflation of the throat. Three different
evenings we tried for flashlights and none show the fully swelled out throat.

MATING

Male (From life, June 21, 1921). Top of head from snout to between
nostrils, over upper lids along back to the vent sudan brown to brussels
brown or cinnamon brown to Prouts brown or snuff brown to bister. Top
of fore hmbs and hind limbs and sides same but less intense. Side of snout
under eye cosse green to light bice green. Same color on back end of upper
eyelid. Spot under tympanum to shoulder insertion, spot over shoulder
insertion, ring in middle of tympanum with dark center, throat and spots on
lower jaw bright chalcedony yellow. The throat reticulated or spotted black
and bright chalcedony yellow. Rim of lower and upper jaw black, the former
with three or four spots on each side. Tympanum same color as back with
some black, then bright chalcedony yellow ring, finally dark center. Under
parts white with black reticulation or deep brownish drab on under parts of
hind legs. Some bright chalcedony yellow on front and inner side of fore limb.
Fore limb with a black stripe continuous from lower jaw to arm insertion on
to the fore arm. Fore arm with rear edge and palm of hand black. Back of
thighs deep brownish drab, black and color of the back.

Iris ring around pupil lemon yellow or part of it light cadmium or orange.
Iris background black but so covered with orange or cadmium yellow spots
as to obscure most of the black.

Female (From life, July 5, 1921). Top of head slightly lighter than back,
near Prouts brown; rest of back and back of tympanum mummy brown or
darker. Or it may be head olive brown and the rest of dorsum clove brown.

358 Frogs of the Okefinokee Swamp

Tympanum like back. No different rings like the male. Spot from arm in-
sertion under eye and forward obscure in part, pale ochraceous buff or light
buff. Under parts with deep brown drab reticulations and cloudings. Spots
onlower jawpalechalecdony yellow. For remainder of description see the male.

Two specimens (From life August i8, 1922). A female dark olive buff
uniform above. The smaller specimen wood brown to avellaneous and uni-
form above. In both back of eye and side of head and over angle of mouth
apple green. Bars on hind legs absent in this female; scanty in smaller
specimen.

Structural differences. Enlarged thumbs and tympana in male; rings of
color in male tympana; throat with bright chalcedony yellow.

"In 1912 we had 14 adults of which twelve were males. Male specimens
of 59-65 mm. show ear drums well developed and this sexual difference may
possibly appear in even smaller specimens. All the males have the forefingers
more or less swollen ; in the two taken after July 15 it is least developed and
the three with it most highly developed were taken June i and 12, 19 12. A
female taken June 5, 191 2 was spent and another female taken May 2 8- July
15, 191 2, is with ripe eggs. These ovarian eggs are 1.2- 1.5 mm. in diameter
and this female has no more than 1,000 eggs to lay. In size she is only 67.5
mm. in length, much more slender and smaller than the average of our northern
green frog females which have from 3,500-4,500 or more eggs to lay."

Duration. In general mating with an individual male is of short duration.
But the species in the northern states breeds for a long period, last of May to
August. In the south it is one of the last to begin breeding and doubtless is
not one of the forms which breeds most of the year. There are one or two
species in the south which might breed almost every month in the year
(except one or two months), but in the south the green frog is not in this class.

Night or day. In general this shy species in the Okefinokee Swamp mates
or begins mating mainly at night. Sometimes in the dark shades of cypress
ponds or "bays" it mates by day.

Amplexation — {Normal, abnormal). Normally it is pectoral in this
species. The first amplexation for this species I ever recorded at Ithaca was
more Hyla-like than Rana-like. The first record for Okefinokee Swamp is of
the same sort. A pair mated in laboratory on Billy's Island, July 4, 192 1,
were more axillary in embrace than pectoral. With only these two examples
one might go astray. The males, however, almost invariably seize the females
in pectoral fashion.

OVULATION

Habitat. They seek small, permanent ponds in the hammocks, the
island edges of cypress ponds and cypress bays in the shade. Under Rana
grylio we noted in one cypress pond how Rana grylio males were calling in the
center of the open pond amongst Pontederia cordata, the Ilyla cinerea males
were in the bushes where open pond and wooded fringe of pond meet and the
Rana clamitans males were in the shade of the large trees near the land's edge.
They are greater shade lovers than any other frog of the swamp region unless
it be Rana heckscheri.

Plate XXXVI

Green frog (Rarut chmitans)

1 . Egg mass, Billy Id., Ga. July 5, 192 1.

2. Eggs, stained, Chesser Id., Ga. July 16, 1922. x 1.2.

3. Tadpole with four legs, Chesser Id., Ga. August i, 1922. x 1.2.

4. Tadpole with short tail, Chesser Id., Ga. August 11, 1922. x 1.2.

5. Transformed frog, south of Spanish Creek, Folkston, Ga. July 29, 1922. x 1.2.

6. Adult female and transformed Eana clamitans, Billy Id., Ga. May 18, 192 1. x 0.3.

7. Transformed R. clnnuians and R. grylio compared, Billy Id., Ga. April 28, 192 1.

Lateral aspect, x 0.4.

8. Transformed R. dantitarifi antl R. grylio compared, Billy Id., Ga. April 28, 192 1.

Ventral aspect, x 0.4.

9. Male croaking, Billy Id., Ga. May 24, 192 1. Flashlight.
10. Male, Billy Id., Ga. April 29, 1921. Ventral aspect, x 0.4.

Rana clamitans 359

In 1922 we found different types of habitats in addition to the above rec-
ords. In Civil War times at Camp Pinckney the Confederate Army dug five or
six large pits. They were going to hide turpentine in barrels in them if need
be. These ponds are now filled with water. In these ponds, some of which are
very much shaded, were several green frogs. The surface of the water was one
continuous mat of fine green vegetation. On July 24 in one of these ponds
we found three fresh masses attached to the base of large grass (4 feet high)
in the middle of the pond. On the road south of Folkston and south of Spanish
Creek is a moist woods, on the side a high sandy ridge. Here in very small
sphagnaceous spring ponds beside the road we found eggs on the moss in
the water.

On July 9, 1922, at Thompson's Landing in a cut off pool left by high
water of St. Mary's River we "found a male R. clamitans at the edge of this
pool under some overhanging rootlets to which a film of eggs was attached.
They are almost hatched. These are not all at the surface. Rains have raised
the level of the pond. These wooded banks of the St. Mary's River have
considerable shade."

Period. In the north the extreme dates for ovulation have been May 23
and August 10. The period of greatest ovulation has been in June and first
of July.

In 191 2 our data for Okefinokee Swamp is: "On June 2, 19 12, on Billy's
Island near Billy's Lake we found in ponds at the edge of the island some green
frog eggs. On the following day in a shallow pond where the water was i
foot deep more were recorded. On June 9, we found another mass and by
June 12 these eggs were hatching. Finally on June 12, 1912, on the north
edge of Lee's clearing we found a pond 15 x 10 feet and 3x4 feet deep in
places. Here we took three male R. clamitans. In the pond were Gambusia,
narrow-mouthed toad, meadow frog and toad tadpoles. Two masses of eggs
were found, one fresh and the other hatching. The masses were films on the
surface of the water as with our northern forms."

In 192 1 the first eggs found came July 5 and the last record July 15. In
1922 we found eggs at Thompson's Landing, St. Mary's River July 9, south
of Spanish Creek July 15 and at Camp Pinckney (near Folkston), St. Mary's
River July 24. In both years July seems the climax month. We thus have a
recorded period from June 2- July 24 for the three years of observation.

Temperature and humidity. In 192 1 on July 3 we had very hard rains in
the swamp; on the 4th, Waycross had an inch. Humid weather preceded the
high period of ovulation July 4-5. No minima went below 69° or 70° and
maxima hovered from 88°-92°. On July 15 minima ran from 69°-72° and
rain occurred throughout the Okefinokee area.

In 1922 the day before Julygthe minima were 6t°--]2° the maxima 82°-93°
and precipitation .25 at St. George, July 9. Our records for July 8 at Chesser
Island were temperatures 92° at 2 p.m., shower at 5:30 p.m. and 73° about
midnight July 8-9, and eggs found July 9. On July 15 and the day preceding
ovulation at St. George, Ga., minima are 66°-68'', maxima 94°-96°, our
lowest at Chesser Island 68° and highest 88° but no rain in either place. On

360 Frogs of the Okefinokee Swamp

July 24 the eggs were found at Camp Pinckney. There were no rains at
Chesser Island, Hilliard, St. George that day or before. Temperatures were
minima 69°-7i°, maxima 9o°-95°. More often I believe precipitation or con-
siderable humidity precede laying, but the above data shows laying without
previous precipitation. The minima range from 66°-73°. Our conclusions for
Ithaca were that the species begins breeding normaliv at 6 5 "-7 4°.

Egg-laying process. Every egg mass recorded m 1912, 192 1 and 1922 was
a continuous film implying that the pair do not move about during oviposition.
In some cases little packets are found scattered on the water amongst float-
ing vegetation. We have assumed that currents, floods, etc., have broken the
original mass in these cases, yet it is possible an occasional pair may move
about during ovulation (e.g., as in Hyla versicolor).

EGGS

Attachment. In 191 2 the film taken June 9 was amongst some pine needles
at the edge of the pond in a hammock and the other masses were about the
edges and attached to growing vegetation. On July 5, 1921, we found at
least 5 masses in all, four in one pond and another in a large pond. "The first
was surrounded by vegetation and grass. A Myrica was overhanging. An-
other mass was at the base of a gum tree in small weeds. The third mass was
in shallow overflow water amongst some growing weeds. The fourth mass
was also amongst weeds and more or less broken up into packets. The fine
mass was attached to the base of a gum tree. There were several male Rana
clamitans around. The masses at a little distance can be easily seen because
of a vegetative scum or oily scum or air bubbles amongst the eggs. The mass
often looks blackish and is sharply outlined on the surface as with films of
Hyla femoralis eggs."

Egg mass. In 191 2 we recorded only the film form for this species in
Okefinokee Swamp. On July 5, 192 1, we found one mass 6 x 12 inches in
diameter. Seldom does either diameter reach a foot. 'Tt is on the surface
of the water, one continuous film, each egg not appearing to be separate in
the film as in Gastrophryne eggs." We found another mass 9 x 10 inches
in diameter and another mass 10x10 inches. In all there were four masses in
one small pond. In general the green frogs of Okefinokee seem smaller than
some of the north and the egg complements are smaller. One complement of
July 15, 1932, we counted. "The total count of the eggs was 1451. This was a
small complement. I hold the complements here (Okefinokee) get twice as large."

Egg description. In 19 14 we described the eggs of Rana clamitans at
Ithaca as follows: "The eggs have white vegetative and black animal poles.
The outer envelopes range from 5.6 to 6.0 mm. in diameter, 5.7 mm. being
the average, 6.0 mm. the mode. The middle envelope may be other than
spherical and often has an elliptical form; it varies from 2.8-4.0 mm. in dia-
meter, the average is 3.3 mm., the mode 3.0 mm. The vitellus ranges from
1.2 to 1.7 mm., the average 1.4 mm., the mode 1.2 mm."

In 191 2 amongst some of the Okefinokee green frogs we noted some eggs
with elliptical inner envelopes.

Rana clamitans 361

In 192 1 some of the first eggs of July 5 were as follows: Vitellus of one
1.4 mm., two more each 1.6 mm.; vitelline membrane of one vitellus 1.6 mm.
is 2.2 mm. in diameter, the vitelline membrane may be quite remote from the
vitellus, sometimes as development proceeds the membrane shows more dis-
tinct than the inner envelope; inner envelope usually pyriform, eUiptic or
sometimes circular, inner envelopes of 2.6, 2.8, 3.0, 3.2, 3.4 mm., most of them
at least 3.0 mm. or more. The outer envelope is seldom distinct if present
but probably becomes a part of the film or jelly matrix of the mass. Later
the same day we make this notation in regard to some other R. clamitans
eggs: "These eggs are fresh. Others described are not so fresh. The inner
envelope is round, 3.2-3.6 mm.; the vitelli 1.2, 1.2, 1.3, 1.4 mm.; vitellus
black and white."

Dangers. None of the Ranas of the swamp lay in such precarious places
as the green frog. It chooses the edges where water levels change most
quickly. I beheve in my experience I have found proportionately more
spoiled R. clamitans masses than with any other species. On July 5, 1922, in
one pool the water reached two inches in depth and the eggs were about at the
hatching stage. Many a time we found the green frog tadpoles in cut off
shallow pools of cypress ponds. In our own pond (at Ithaca) we find that the
frogs, particularly males and sometimes females, are easier of capture at
breeding time. The toll of frogs from water snakes (Natrix s. sipedon) is
considerable. We have seen water snakes seize more green frogs than any
other species of frog and have often made these snakes release their quarry.

HATCHING PERIOD
In 19 1 2 one mass of June q hatched in three days.

MATURE TADPOLE

Color description from life ( Non Ridgway) . Background of back very dark
and covered with very fine yellow spots, the whole consequently having an
olive green color with numerous distinct dark spots. Belly deep cream color
without decided iridescence. The throat and sides are mottled with dark
green. A slight coppery iridescence on the venter is more decided on the sides
and on the tail. The tail appears green, mottled with brown; it is covered
with fine yellow spots like the back.

General appearance. Tadpole large, not deep bodied. Tail fairly elon-
gate, tip acute. Dorsal crest not so wide as musculature, extending forward
on body sHghtly ahead of the vertical of the ends of the hind legs. Spiracle
sinistral, just visible from dorsal aspect, directed obliquely upwards and back-
wards. Spiracle below lateral axis. Spiracular opening elliptical and plainly
visible as such. Muciferous crypts very distinct in life, indistinct in most
preserved specimens. Eye on or above lateral axis and near lateral outHne
in dorsal aspect than mid dorsal line. Anus dextral, opening on level with
edge of ventral crest.

Mouth parts. Edge of upper labium fringed with teeth and about equal
to upper horny beak in length as in R. grylio. In either corner, beneath this

362 Frogs of the Okefinokee Swamp

fringe is a very short row (sometimes absent) which is from 2/1 5-1 /16 the
upper fringe. The ends of the lateral row not extending beyond end of upper
fringe. Median space between lateral rows six to eleven times the length of
either row. The first lower labial row slightly longer or equal to horny beaks
in length, and sometimes divided in the middle. The second row almost
equal to first. The third row quite short not nearly as long as in R. catesheiana
usually almost 1/2 of the first row not 3/4 or 4/5 as in R. catesbeiana, or 2/3 as
in R. grylio. It is contained one and one-half times in the horny beak and as
much shorter than the single row of lower papillae. On the side of the labium
the inner papillae (inside outer row) extend mesially beyond and beneath the
second lower labial row of teeth, but not to third row (more like R. grylio).

Measurements. Length of body (12.2-27.8 mm.) 1.5-2.1 in tail (18.4-57
mm.), average 1.75. Width (9.8-21 mm.) of body i. 25-1. 7 in its own length
(12.2-27.8 mm.) average 1.47. Depth of body 1.13-1.32 in body width,
average 1.22. Depth of body 1.9-2.3 in length of body, average 2.14. Depth
of tail (7.4-16.6 mm.) 2.5-3.7 ^^ length of tail (18.4-57 mm.) average 3.1.
Muscular part (3.8-10.4 mm.) i. 35-1. 95 in depth of tail (7.4-16.6 mm.).
Spiracle i. 35-1. 8 nearer base of hind legs or vent region. (5.4-12.4 mm.) than
the tip of the snout (8.2-19 mm.). Spiracle i. 06-1. 38 nearer eye (4.4-9.4 mm.)
than base of hind legs or vent (5.4-12.4 mm.) average 1.24. Eye distinctly
nearer to tip of snout (3.6-7.8 mm.) than to spiracle (4.4-9.6 mm.). Nostril
decidedly nearer eye (1.4-36 mm.) than snout (2.2-4.8 mm.). Mouth (2.6-
4.8 mm.) .8-1.35 larger than internasal space (2.8-4.8 mm.) average 1.06.
Mouth contained 1.3-1.8, average 1.5 in interorbital distance (3.8-8.2 mm.)
Internasal space contained in interorbital space 1.25-2.05, average 1.6.

The dimensions of the largest tadpole are :

Total length

64.8

Spiracle to vent

12 . 2

Body length

27.8

Spiracle to eye

9.6

Body depth

17.2

Eye to snout

7-4

Body width

20.6

Eye to nostril

3-6

Tail length

37-0

Nostril to snout

4.0

Tail depth

15 -6

Mouth

4.6

Musculature of tail

10. 0

Interorbital distance

8.0

Spiracle to snout

19.0

Internasal distance

4.8

General remarks. On July 9, 1922, at Thompson's Landing, St. Mary's
River we found in a pond sotne tadpoles almost white in appearance yet they
were not albinos. They were in a little pool with chalky white water and were
colored accordingly.

LARVAL PERIOD

Normally I believe this species spends one winter as a tadpole. In 192 1
it was difficult to find tadpoles inside the swamp after May 15. In 1922 we
found more in shaded ponds on the outside east of the swamp. On July 15
in one sphagneous pool where the species was laying we found a few tadpoles
nearing transformation. On August 11 at Camp Pinckney we found quite a
number of large tadpoles in five deeply shaded pools. But even on the outside

Rana clamitans 363

mature tadpoles after June i are scarce. I am aware of Swingle's striking-
experiments with hastening transformation in this species, and am also fully
conscious this species has a longer growing period each year in the south than
in the north, but I nevertheless believe one year its normal tadpole life in the
Okefinokee region.

TRANSFORMATION

Period. In 191 2 the writer was in the swamp from May 28- July 4 and
the species was almost finished transforming by the later date. In 192 1 we
have definite records for April 27 and May 14. Our collections show 14 trans-
formed or recently transformed from April-August, 192 1.

In 1922 we found on July 15 and July 17 some tadpoles with large hind
legs but they would not have transformed before August i or 10. One trans-
formed July 28. Most of the transformation comes in the spring and early
summer. Transformation stages were hard for us to find in the Okefinokee
Swamp after June 15 or July i.

Size. At Ithaca the range of transformation size is generally 28-38 mm.
but many of those of the Okefinokee are smaller. In 191 2 on June 9 we found
some tadpoles one of which would have transformed in a week or ten days. It
was 23 mm. long without the tail. From May 28 to September a series of
frogs just transformed measured 25.5-33.5 rnni.

In 192 1 on April 27 we visited a large cypress pond west of camp. In little
cut-off pools where there was little vegetation and mainly pine needles we
found two transformed green frogs, 23 and 25 mm. in length. Fourteen more
specimens collected from April-August, 192 1, measure 20, 21, 22, 22, 22, 22,
23, 24, 24, 24, 24, 25, 25, 28 mm. respectively. On May 14, 192 1, we took two
transformed frogs 21 mm. and 24 mm. in length.

In 1922 on July 17 in one pond we found a few tadpoles nearing trans-
formation. One transformed July 29, 1922, at the small size of 20.5 mm. We
have a range of 20.5-33.5 mm., or probably with an average near 25 mm. or
at least 7 mm. smaller than the average transformed green frog of the North-
eastern States.

General remarks. Possibly inasmuch as the adult of this species in the
Okefinokee region never attains the size of the northern representatives, we
have a smaller size at transformation as well. In any event both adult and
transformation sizes are smaller. The transformed frogs are easy to tell be-
cause of their prominent heavily spotted breasts. These numerous small
spots are grayish white.

GROWTH

Of 27 adults, 4 were from 52-58 mm., 16 were from 60-69 mm., and 7
from 70-79 mm. It must be rarely that Okefinokee representatives of this
species exceed 80 mm. in body length, yet northern forms may reach 100 mm.
The mean range of adults is between 60-70 mm. On June 12, 1912, we took
male frogs 52, 66, 67, 70, 72 mm. in length or on June 5, 1912, one 29 mm. and
a female 58 mm. Between May 28- June 25, 191 2, we found small frogs from
25-33 mm. in length. In the same period we took adults 52-75 mm. but none
in the gap of 34-51 mm. In 192 1 we secured only two in this intermediate

364 Frogs of the Okefinokee Swamp

category, one 43 mm. June 24, 1921, and one 40 mm. July 5, 1921. On this
later date we secured one 40, 65, 69, 7 1 mm. respectively. We took two young
of 22 and 32 mm. from April 22-May 23, 1921, 14 young from 20-28 April-
August, 1921, two young 28, 32 mm. June 20, 1921, and one 33 mm. July 26,
192 1 — in other words 20-33 mm. in range. This gives a clue in April and
May of 192 1 of transformed frogs 20-33; others 40-43 mm.; others 56-62
mm.; others 65-72 mm., or in June and July, 1922, we took frogs 20.5 mm.
at transformation, others 59 and 60 mm., and others 61-77 mm. The growth
stages then seem 20-33 mm. at transformation; 33-43 mm. the first year's
growth; 43-52 mm.; 52-63 mm.; 64-77 mm. Possibly these four presumptive
years based on measurements of frogs taken at the same period should be
shortened to three years.

FOOD
We have made no study of the food of the specimens we captured.

ENEMIES

In 19 1 2, the only enemy of which we had positive evidence is the moccasin.
One moccasin taken June 22, 19 12, had a two-thirds grown R. clamitans in its
stomach and another secured in June, 19 12, had a full grown frog.

In 1922 an amusing episode reveals another unusual enemy. On June 27,
at Camp Pinckney (near Folkston), Ga., on the St. Mary's River were some
big ponds (formerly dug by Confederate Army to hide turpentine in barrels).
Here we found several green frogs. "I captured one and started for another
pond. Halfway I lost the frog and began to chase it but an old hen ran in and
seized it ahead of me. It ran off with the prey and ate it. The frog was two-
thirds grown." See ovulation (dangers) for discussion of water snake as an
enemy.

AUTUMNAL DISAPPEARANCE

Our latest record is for September 5, 1922.

BIBLIOGRAPHY

See General Bibliography under synonymy Boulenger, G. A. (Proc. Am. Acad. Arts &

Science, LV, 1920, p. 425).
1914 See previous notes, A. H. Wright, 1914, Carnegie Inst. Publ. No. 197, pp. 70-76;

1920, Rept. U. S. Bur. Fisheries, Doc. 888.
1914. Overton, Frank. Brooklyn Mus. Quarterly, March, 1914, p. 37.

Rana grylio Stejneger

(PI. Ill, Fig. 8; VIII, Fig. 4; XI, Figs. 5, 6; XIV, Fig. 5; XVI, Figs. 3, 6; XVII; XXXVL
Fig. 8; XXXVII; Text Figs. 1,15)

COMMON NAMES
Southern Bull-frog. Joe Brown Frog. Pig-frog. Lake Frog. Bull Frog.

RANGE

Check list. Rana grylio Stejneger. "Range: Southern Mississippi to
to peninsular Florida. Type Locality: Bay St. Louis, Mississippi." — Stejneger
& Barbour (Check List, 19 17, p. 38).

Rana grylio 365

Supplementary records. Dr. Stejneger (1901) had it from Kissimee,
Pensacola, Florida and Bay St. Louis, Mississippi, and gave its range as
"Florida and Gulf Coast west to Mississippi." In 1906 Miss Dickerson re-
ported it from Ozona, Florida. In 191 2 the author captured several specimens
and saw many more in Okefinokee Swamp, Georgia. From 1914-1922
Deckert reported it from Jacksonville, Royal Palm Hammock, Musa Isle,
Miami River, Florida. In 1916 T. Barbour reported it in some of the Florida
lakes and in 1920 he notes it near Palm Beach, Florida. In 1922 Loding in-
cluded this form with the comment "Alabama records: none." However,
the distinctive note of this frog has been heard by the writer and others in
Mobile County. In 191 7 our transcontinental party of 13 individuals (Am.
Naturalist, April, May, June 19 19) camped near Theodore, Alabama. Prof.
J. C. Bradley remarked "Across the road from our camp, some peculiar
alligator-like grunts proved to be emanating from the southern bull-frog,
Rajia grylio. This species was described only about fifteen years ago." We
captured and photographed two or three of these adults. In 192 1 and 1922
we found them through the Okefinokee Swamp. In two different articles
Viosca in Louisiana (1923), noted their presence throughout "the entire
southern portion of this state," a westward extension of its range. Earlier in
19 1 8 Viosca writes "During the last year I have made positive identification
of this species in many localities in the southern part of Louisiana." Boulenger
(1920) secured specimens from New Orleans and Plaquemine, Louisiana. It
is now known from Georgia, Florida, Alabama, Mississippi and Louisiana.

Inasmuch as we called this species a Sabalian species in 1924 (Ecology,
January 1926, Vol. VII, p. 82) we would expect it to extend to South Carolina
or Wilmington, N. C, and records from eastern coastal Texas are to be
expected.

Local Okefinokee records. This is one of the characteristic notes of all the
water lily prairies of the swamp and the open cypress ponds. We also re-
corded it in the grassy or weedy cypress bays, along the river swamps of the
Suwannee and St. Mary's Rivers, along the floating vegetation of the edges
of water courses or lakes in the wooded swamp. It is decidedly aquatic and
breeds in its accustomed habitat of the rest of the year.

GENERAL APPEARANCE

This is a southern bull-frog of the Sabalian or Humid division of the Lower
Austral Zone with shorter snout, narrower head and shorter fourth toe. In
coloration it is an overgrown Rana virgatipes without two dorsal stripes, and
is a beautiful frog.

COLORATION OF SPIRIT SPECIMENS (19 1 2)

General back color is olive citrine or brownish olive, bone brown or blackish
brown all over. In small specimens there is hardly a dark spot, but in the
olive forms the dorsum from the tympanum backwards is covered with
prominent black blotches with those on each side from tympanum to groin,
more or less merged. On the forelimbs of all the specimens there are no spots

366 Frogs of the Okefinokee Sivamp

or bars. On the hind Hmbs of the younger forms and in the ohve colored group
there may be three cross bars on the front half of the femur, three or four
bands across the tibia, and three or four on the tarsus and foot. In the rich
or deep brown specimens the dark spots obscurely show. Occasionally in
these specimens the black body color or sometimes a black color almost
wholly covers the upper two-thirds of the posterior faces of the thighs. In
all the specimens the body color covers one-half to two-thirds of the posterior
faces of the thighs. Occasionallj'' the body color is replaced by black. Beneath
this area of body color comes a longitudinal yellowish white or yellowish stripe
followed by a prominent dark, black or brown band. Sometimes this is
succeeded by another yellowish line followed by a wonderfully brilliant
mottling of brown and yellow. In almost all of the largest forms where
mottling on under surface of the legs is prominent these yellowish lines are
yet indicated by two broken up longitudinal rows of yellow spots. In some
the second yellow line is absent. The smaller specimens of females have the
underparts except the thighs immaculate. One large male has the whole
under parts the most striking black or brown and yellow we have seen in a
North American frog. On the posterior face of the tarsus and foot is a
prominent dark band with a yellowish line below it on the tarsus. This colora-
tion shows best in young forms. These specimens have the forearms unicolor
above. The sides of some of the specimens are unspotted while others have
some mottling of black chocolate brown and yellow like the venter. Three
large males have the under surface of the leg speckled, mottled or reticulated.
On either side of the breast and in front of the arm insertion, the dark body
color, gray or brown extends as a special area almost or actually to the middle
line. Most of the specimens have the whole upper head unstriped or un-
spotted. The tip of the snout is more or less white. The web is dusky, in
adults very clouded; in young, clear with numerous spots or mottlings.

STRUCTURAL CHARACTERS (WRITTEN IN 1 91 2)
Head smooth, other parts minutely mammillate and pitted, sometimes
round on the sides, throat, and lower belly; snout more pointed than in Rana
catesbeiana; eyes elevated, large; eyelid rough; nostrils equidistant from eye
to tip of snout or slightly nearer the snout ; a prominent fold of skin from the
eye over the tympanum to above the shoulders; no lateral fold; groove down
middle of back halfway to the vent or sometimes absent; first, second and
fourth fingers about equal, long; inner metatarsal tubercle small; no outer
tubercle; subarticular tubercles very small; Stejneger ('02) found the per-
centages of toes measurements in Rana grijlio and Rana catesbeiana to be as
follows: in Rana grylio the third toe in the length of the fourth toe is .81-.84;
second toe in the fourth toe, .55-.61; first toe, .34-.39; in Rana catesbeiana,
third toe, .70-.76; second toe, .47-.51; first toe, .27-.33. In some of our
specimens of Rana grylio the proportional length of the third toe in the fourth
is even greater than Stejneger found, being sometimes .85 or .86 while the
second toe may be as much as .65 and the first toe .43. The head measured
at the tympanum is perceptibly narrower than R. catesbeiana; vomerine teeth

Plate XXXVII

Southern bullfrog (Rana grylio)

1. Male, Billy Id., Ga. April 29, 192 1. Ventral aspect, x 0.4.

2. Female, Billy Id., Ga. June 22, 1921. Ventral aspect, x 0.4.

3. Transforming frog. Billy Id., Ga. April 26, 1921. Lateroventral aspect, x 0.4.

4. Male croaking, Long pond, Billy Id., Ga. May 19, 192 1. Flashlight.

5. Mature tadpole, Billy Id., Ga. August 4, 1922. x i.o.

6. Tadpole with four legs, Billy Id., Ga. .June 5, 192 1. x 0.4.

7. Transformed frog, Billy Id., Ga. April 26, 192 1. x 0.4.

8. Site of egg mass in cypress glade west of Minne Lake, Ga. June 10, 1921.

9. Bullfrog pond looking S. S. W. Billy Id., Ga. June 16, 192 1.

Rana grylio 367

close together often obliquely placed; tongue with moderate forks like Rana
catesheiana or with extreme elongate thin attenuate horns; no dorso-lateral
glandular fold; webbing of hind feet less emarginate than in Rana catesheiana.

MEASUREMENTS

(1912-1914)
The measurements of 7 specimens follow. The total length reaches from
53-101 mm.; average 82.3 mm.; the head is 21-39 mm., average 31.6 mm.,
usually in males larger than the width of the head, in females equal to or less
than the width of the head; the length of the head is contained in the length
of the body 2.3-2.8 times, average 2.5; width of the head, 21.5-40.5 mm.,
average 30.6 mm.; snout 8-15 mm., average 11.5 mm., greater than eye in
all, in males usually less than (rarely equal to) the tympanum, in females
greater than the tympanum; eye 7-1 1 mm., average 9.6 mm. less than snout,
less than the tympanum in males, equal to or greater than tympanum in
females; tympanum 5.5-17 mm., average 10.9 mm., large in males, moderate
in females; interorbital distance 3.5-6 mm., average 5 mm., from 1-2 times
in upper eyelid; femur 25-51 mm., average 38 mm., equal to tibia; tibia 25.5-
50 mm., average 39 mm., slightly less than rest of the foot without tarsus;
tarsus 15.5-29 mm., average 22.5 mm.; rest of foot 28-34 mm., average 22.5
mm.; rest of foot 28-54 mm., average 41.8 mm., anterior limb from axilla
30-52,5 mm., average 41.6 mm., more than femur or tibia, less than equal to
or more than rest of hind foot; posterior limb from groin 77,5-141 mm.,
average 114,2 mm., 1.3-1.5 times the total length; posterior limb to heel
reaches the front of the eye or even to the nostril.

(Recent Material)

Head to angle of mouth 1.02 (50 mm.)-i.o2 (51 mm.)-i,i3 (82 mm. d^)
-1.08 (95 mm. cf) in width of head; head to rear of tympanum .95-.975-.94-
.95 in width of head; head to angle of mouth 3.37-2.68-2.82-2.71 in length;
head to rear of tympanum 2.8-2.55-2.34-2.375 in length; snout i.o-i,i2-.92-
i.o in fourth finger; 1.06-1.12-1.15-1.06 in first toe; eye 1.14-1.14-1.44-1.6
in snout; eye .70-. 85-1. 44-1. 6 in tympanum; eye 1.14-1.29-1,55-1.5 in first
finger; tympanum 2.4-1.66-1.2-.93 in intertympanic width; i. 6-1. 33-1. o-i.o
in snout; internasal width .88-.875-.857-,85 in upper eyelid width; inter-
orbital width 1,33-1.16-1. 2-. 86 in upper eyelid width; i. 5-1. 33-1. 4-1.0 in
internasal width; 4.0-3.33-3. 1-2. 14 in intertjTn panic width.

Forelimb: i. 68-1. 7-1. 64-1. 86 in length; 2.6-2,56-2,56-2.8 in hindlimb;
first finger i. 44-1. 33-1. 28-1. 33 in third finger; second finger i. 27-1. 2-1. 1 2-
1.25 in third finger; ,94-.9-.87-.93 in first finger; third finger 1.2 i-i. 08-1. 33-
1.25 in second toe; fourth finger 1.2 i-i.o-i. 25-937 in first toe; 1,64-1,33-
1.51-1.25 in third finger; internasal width 1.77-2. 25-2. 0-2. 14 in first finger;
2,0-2.5-2.3-2.3 in second finger; 2.55-3.0-2.57-2.85 in third finger; 1.55-2.25-
1.71-2,3 in fourth finger.

Hindlimb: length 1.5-1,51-1.56-1,51 in hindlimb; tibia 2.0-2.0-1.95-2.04
in length (snout to vent) 3.0-3.08-3,04-3,13 in hindlimb; 1,15-1,2-1.19-1.18

368 ■ Frogs of the Okefiiiokee Swamp

in forelimb; 1.04-1. 08-1. 14-1. 06 in hind foot; first toe i. 64-1. 44-1. 6-1. 49 in
second toe; 2. 10-2. i i-i. 93-2.0 in third toe; 2.09-2.55-2.93-2.64 in fourth toe;
2.23-2.1 1-2. 33-2. 17 in fifth toe; second toe i. 64-1. 46-1. 2-1. 36 in third toe;
1. 8-1. 76-1. 83-1. 8 in fourth toe; i. 35-1. 46-1. 45-1. 48 in fifth toe; third toe
1. 39-1. 2-1. 5-1. 32 in fourth toe; 1.05-1.0-1.2-1.08 in fifth toe; fourth toe
1.04-1.17-1.09-1.09 in hind foot; 1.0-1.08-.95-1.02 in tibia; 1.16-1.3-1.13-
1.13 in foreUmb; fifth toe 1.31-1.2-1.25-1.21 in fourth toe; internasal width
1. 88-2. 25-2. 14-2. 43 in first toe; 3-i-3-25-3-43"3-57 in second toe; 4.0-4.75-
4.14-4.85 in third toe; 5-55-5-75-6.3-6-43 in fourth toe; 4.02-4.75-5.0-5.3 in
fifth toe.

HABITAT

This is ideally a frog of the prairie or open centers of cypress ponds which
are an extension or separation from the prairie types themselves. Waterlihes,
hardheads (Xyris) neverwets (Orontium), wampee (Peltandra), watershield
(Brasenia), bladderworts ( Utricularia), floating heart ( Nymphoides) prevails.
Where this habitat approaches islands, bays, or a circle of trees, maiden cane
(Panicum), fern (Woodwardia) and duck grass (Rhynchophora) occur. Where
this latter assemblage of plants occur in bays, etc., Rana grylio may be or
where some of it may be along water courses.

FIRST APPEARANCE

The earliest definite date of their reappearance is Dr. Barbour's note. At
Palm Beach, Florida, "Rana grylio reappeared in 1920 on March 17 after not
being seen all winter." We have not been afield in its range except in April
to September in which period it was active in 1912, 1917, 1921 and 1922.
Deckert in observations from February 23-December 8, 1920, first heard this
species grunting April 4, at Musa Isle, Dade Co., Florida.

GENERAL HABITS

Mr. Robert Ridgway found them "so excessively shy and wary that he
had failed to secure any specimens" for Dr. Stejneger. "Requests were sent
to various correspondents to catch and forward specimens, but they were
equally unsuccessful." Stejneger 1901 (p. 2 1 1), also Dr. E. A. Mearns reported
them very difficult of capture. In 191 4 Miss Dickerson says "It seldom leaps;
its strong instincts when frightened are to dive and hide." Deckert found it
in 1911-1912 (1914, Copeia, No. 3) at Jacksonville, Florida "not rare, but
very shy." Later he pronounces that "It is one of the shyest of all frogs,
usually floating in water two or three feet deep, under branches of a floating
dead tree, or among other debns, so that it is almost impossible to get within
capturing distance without alarming it. Its size is 4 to 5 inches from snout
to vent." In 192 1 he writes "No effort was made to take specimens, as it is
impossible to secure living material during the daytime, and without a boat,
the species being so thoroughly aquatic and exceedingly shy." In (1920) Dr.
Barbour calls it "very shy and wholly aquatic." Viosca in 1918 (p. 161)
writes "It appears to be confined to the alluvial section of Louisiana, where it

Rana grylio 369

frequents the freshwater marshes, and is especially abundant in the many
lakes and lagoons of these regions. The individuals are very shy, and are
seldom seen by daylight, as they hide among the aquatic vegetation of the
lakes, and when closely approached quietly take to the water and hide in the
mud — not diving with the splashing retreat of the common bullfrog. They
are entirely aquatic and are rarely caught on the land."

In 191 2 the following notes on their habits and habitat were made: "One
will never forget his first capture of the 'Swamp bullfrog'. The prominent
black or brown and yellow reticulations of the under parts are very conspicu-
ous. Sometimes we succeeded in kiUing them with an oar or pole while
pushing through the bonnets and occasionally the members of the party
captured them when visiting set trap lanterns for insects."

On April 25, 192 1, "we went with flashlight after Rana grylio. On the
vegetative carpet and lily pads were untold numbers of Acris, on the lily
pads and on the bushes Hyla cinerea, and amongst the pickerel weeds different
sizes of Rana grylio. In all, three of us secured only 6 adults and one trans-
formed specimen. Later I found that they could be picked up rather easily
with a flashlight. On May 6, Dave Lee went out on Billy's Lake at night
with a torch to catch small fish for bait. With a small dip net he would scoop
in front of a Rana grylio and usually catch them."

On May 10, 192 1, we found one cypress pond with plenty of all stages of
Rana grylio. Our comment is that they are "easy of capture at night except
for adults and these even can be taken." On May 11 we found plenty of R.
grylios at night along the edges of Billy's Lake. "They usually were at the
edge near the bushes or under them or amongst brush or in maiden cane.
Usually when one tried to photo them they were too much surrounded with
vegetation or sticks. We could approach closely but in clearing away brush
for the flashlight photo we would scare them. Often we would hear them go
skipping along across the lake or along the edges like a Rana catesbeiana."
As with most shy frogs they can be captured if females are around. In some
cases the females led the way to where a male might be. Where we succeeded
in flashlight ing a croaking male we were quite certain females were near
because of eggs found there soon afterwards. On June 20 we found another
method of capture. We never dreamed it would work for Rana grylio. We
would pull in hurriedly, masses of floating maiden cane and in the corner at
the bottom of the boat all kinds of life would drop. On this night we caught
2 Rana grylios, one a fine ripe female. All in all our main reliance was to
wade about at night on the prairies or in open cypress and to catch them by
hand. One must, however, keep his bearings.

Anent this species it might be interesting to include some comments of
Percy Viosca, Jr., in a letter to Dr. H. F. Moore, U. S. Bureau of Fisheries.
On July II, 1923, he writes: "It might also interest you to know that Rana
grylio is abundant in this State, but we have found it is a thoroughly aquatic
species and not very hardy in captivity, and would not be suitable for frog
culture even though it is a fine flavored species and we believe much superior
to flesh of the catesbeiana. It would be only suitable for planting in shallow

370 Frogs of the Okefinokee Swatnp

lakes of the lagoon type, but once established it would be difficult to collect,
except by experienced night hunters. It would be a species very difficult to ship.
Although we have shipped them successfully, the frogs need individual care."

VOICE

In I go I, Dr. Stejneger remarked that "A few years ago Mr. Robert
Ridgway, returning from a collecting trip in Southern Florida assured me
that the bullfrog there had such a peculiar voice that he could scarcely believe
it to be the same species as the one found elsewhere in the United States."
This was the genesis of Dr. Stejneger's interest in this new form. Later Dr.
E. A. Mearns sent some from Kissimee, Florida "with the statement that
their voice was entirely different from that of northern bullfrogs, resembling
the grunt of a herd of pigs."

In 1906 Miss Dickerson held that "The frog might be unknown now, if it
had not been for the persistent reports of its voice. The sounds produced are
said to resemble the grunting of a herd of pigs, thus differing entirely from
the familiar bass notes of the common bullfrog."

"Rana grylis Stjr.," Deckert, 1914, writes "is called the 'Pig-frog' owing
to its loud grunting call, repeated three of four times. . . . The call is heard
at any time, day or night, from deeply wooded bayous, oftenest in March,
April and May." "In May 1922 we heard it at Royal Palm Hammock, Dade
Co., Florida."

To Dr. Barbour in 1920 "Its call, which is heard at night, or on damp days,
resembles the grunting of a pig, and consists of but one sound oft repeated.
Dr. J.C. Bradley in 1919 (June 12) called its note 'some peculiar alligatorlike
grunts'. ... Its note is altogether different from that of our northern species."

In 19 1 8 Viosca held "The voice is entirely different and is much less musi-
cal than that of the common bullfrog. The individual call is a series of grunts,
usually eight uttered in quick succession. When several are calling, the noise
is not unlike that of as many hogs, and 'pig' frog might prove an appropriate
popular name."

In 191 2 we made the following notes on the swamp bullfrog in Okefinokee
Swamp. "From June 13- June 19 we heard several of these within the swamp.
On June 18, 19 12 at the Still Heronry amongst the swampy tangle of button-
bush (Cephalanthus occidentalis) 'hurrah bush' (Leucothoe racemosa) and
leather leaf (Clethra alnifoUa) we heard plenty of southern bullfrogs from 8-1 1
a. m. The water was waist deep and the bushes 8-12 feet above the water.
One of their notes was very much like that of Rana catesbeiana of the north.
Another note frequently given in chorus was mournful, reverberating, prolonged
and deep, not at all like the regular bullfrog chorus note. There were a number
in the chorus and it sounds like one continuous long deep roar. The following
day, June 19, from 7:10-11 a. m. when the sun was very hot we heard several
in the tree-covered overflowed banks of the Suwannee River. On June 13
while travelling through Minne Lake Run we heard the deep, guttural notes
of the 'Swamp Bullfrog' all along the bonnet borders and through the cypress
woods on either side."

Rana grijlio 371

In 191 7 on June 12 two miles from Theodore, Ala., we "heard the croak or
grunt (4 or 5 notes) of the southern bullfrog. Hard to find. Lost the first one.
They are amongst cat briars. Found a young one and later took a large male.
To one member of the party there seems something of the human in its voice.
To another it had an alligator quality. Certainly it differed from the northern
bullfrog."

In 1921 we began hearing R. grylio from the very first of our entrance into
the swamp. On April 23 we heard the first one and it continued a puzzle for
about two days. From April 25 we heard them most commonly from 7 to
8:00 p. m. onward. Several also called during the day. From April 30-May r
I have the note "No Rana grylios heard." "Too cold for R. grylio calls."
From April 23-April 29 our lowest temperatures in early morning (6-7 a. m.)
were 62° and our highest 68°. On April 30-May i it was very cold and the
early morning temperatures were 45°-52°. From May 1-7, 45°-57° obtained
and very few were heard. From May 8-14 a few were heard, low minima 6o°-65°
obtaining at night. From May 16 onward there were plenty of R. grylio
calling in early evening or wonderful choruses at midnight or in early morning
hours before dawn. Minima were from 6-j°--jo° or higher. There after in
June and July we heard them commonly at night and occasionally they were
abundantly heard in the daytime. In general 45°-65° minima are not condu-
cive to calling. From 65° 70° starts calling in numb?r3 and large choruses are
from 7o°-85° or higher. Humidity and rainfall are, however, even more im-
portant than temperature in determining their activities.

The records of 192 1 and 1922 combined give the croaking period as April
23-September 5. Doubtless had we been in the Okefinokee before and after
these dates we would have heard them.

In 192 1 we have the following notes on the call of the southern bullfrog.
On June 25 in mid forenoon "I heard the croak of one frog in the middle of
Long Pond. It sounded like the roar of our northern Rana cafesbeiana.
Heard several normal Rana grylio at 10:30 a. m." On April 25 we record the
call as "two or three grunts of less than a second each. Sometimes there are
more." The R. catesbeiana-like call given June 25 was not in abnormal con-
ditions but on April 25 we carried several in a bag to our camp. Several times
on the way one male uttered a "distress" or "mercy" note not unfike the
familiar "jug-o-rum" of Rana catesheiana. On May 11 we went to Billy's
Lake at night. Along the lake either side were numerous R. grylio croaking
• — just two or three croaks to a male."

In 1920 (p. 21) the author wrote of the call as follows: "In Southern
Alabama he recorded croaking males as not uncommon in overflowed areas
and swamps (Plate IX No. 2) of clear streams, especially if overgrown with a
thick mat of cat briars (Smilax) and arrow arums. In the main their croakings
consist of four or five notes and are wholly unlike the call of the Northern
bullfrog. To some people there is something of the human voice in their call;
to others it sounds like an alligator. If the ventrilocjuial males be in tangles
they are hard to discover. These croaking males may also occur along the
deep wooded overflowed banks of southern rivers."

372 Frogs of the Okefinokee Swamp

Some of the residents call this species the "Joe Brown" frog because of a
fancied resemblance of the grunt to this name. The croak can be heard at a
considerable distance and a midnight chorus might be pronounced "loud."

Mr. Francis Harper describes the note as follows: "Its deep bass note is a
rough, guttural, grunting grro, grro, ... It is usually double (as just repre-
sented), but sometimes single, and occasionally it runs into a series of from
three to ten notes."

The male frog has "internal vocal sacs" says Boulenger. In this regard
it is like Rana damitans and Rana catesbeiana. The green frog inflates its
vocal sac more on the sides below the angle of the mouth and less beneath the
tips of the lower jaw. On the bullfrog it is one swelling below lower jaw. In
the southern bullfrog the inflation is greater than in the other two species.
There is one inflation but on either side of the middle is a subsidiary inflation
giving the internal sac a tripartite appearance (PI. XXXVII, Fig. 4). These
creatures are hard to flashlight at the moment of inflation. The frog is spread
out on the surface of the water amongst the water plants. The call is very short.

Probably Bartram's description is of Rana grijlio when he says that "The
largest frog known in Florida and on the sea coast of Carolina, is about eight
or nine inches in length from the nose to the extremity of the toes; they are a
dusky brown or black colour on the upper side, and their belly or under side
white, spotted and clouded with dusky spots of various size and figure; their
legs and thighs also are variegated with transverse ringlets of dark brown or
black, and are yellow and green about their mouth and lips; they live in wet
swamps and marshes, on the shores of large rivers and lakes; their voice is
loud and hideous, greatly resembling the grunting of swine (our emphasis),
but not near as loud as the voice of the bullfrog of Virginia and Pennsylvania;
neither do they arrive to half their size, the bullfrog being frequently eighteen
inches in length, and their roaring as loud as that of a bull."

MATING

Male {From life, April 25, 1921). Upper parts rich brown with black
spots obscured. Tympanum outer circle deep brown, center green or dark
green. Top of head dark green. Belly creamy. Throat lemon yellow with
dusky fine blotches on underside of fore limbs and hind hmbs. Lower belly
blotched or reticulated with cream and brown. On sides of body and legs
cream replaced by yellow. First finger of male much enlarged. In another
male head deep bottle green. Tympanum center black, middle ring light
brown, outer ring brownish black. Throat almost orange yellow on either side
of the breast. On either side of breast between arm insertions a deep green
area. On rear of femur the yellow extends as a yellow line with brown black
on either side of it. This line golden or orange yellow. More intense than
yellowish or cream of undersides of legs. In some the black spots of the back
are farther apart. Black spots of back surrounded by green and the centers
with bronzy brown. Top of toes brown and yellow. Under side of toes brown.
In half grown male under color cream slight wash of yellow. Femoral stripe
cream not yellow. Ear drum small all brown. Two pectoral patches dusky
not green. (Non Ridgway.)

Rana grylio 373

Female {From life, June 21, 1921). Upper lip from tympanum to snout
medal bronze also on top of head; upper parts olive citrine with a few dark
spots on the posterior back and groin. Upper part of legs saccardo olive with
cross blotches. Throat baryta yellow or better straw yellow. Where dorsal
color reaches light venter, so also on fore and hind legs there is one clear dull
green yellow or light dull green yellow. Venter marguerite yellow, also same
color on top of toes and fingers. Two patches on either side of throat and near
pectoral region, so prominent in the male, is light yellowish olive, not pro-
nounced as in male. No reticulations on venter except for faint reticulations
on under side of hind legs and the slightest indications on the groin region.
Rear of femur with one long band of sea foam yellow with line of quaker drab
below and dark quaker drab or black above. Iris with bright green yellow
ring around pupil. Iris black with lots of bright green yellow spots. Green
yellow outer circle around the black.

Structural differences. The smallest male to show enlarging of tympanum
is 67 mm. or about one year after transformation. All the other of this series
are as follows: None of the other year-olds show secondary male characters
except one at 67 mm. where the tympana are enlarging; in some of the two-
year-olds (73-86 mm.) the tympana are enlarged and half of them have the
thumb discolored. Not until the three old stage (91-102) is reached do they
have the tympana very large and the intertympanic space much reduced.
The thumb is very well developed but never so swollen and stubby as in /?.
virgatipes, R. clamitans or R. catesbeiana. The males have more pointed
narrower heads than the females, are slimmer, have the intertympanic space
much reduced in males, thumbs discolored and somewhat swollen. The
females become larger than the males.

Duration. It doubtless is shorter in this species than in R. sphenocephala
or species which begin ovulation earlier in the season. It is conceivable that
this member of the R. catesbeiana group like the R. catesbeiana might continue
amplexation for a long time. One R. catesbeiana male was recorded mated
with a dead female which doubtless died from a long amplexation. This is
the exception. Normally we would expect R. grylio to be of a short amplexing
group.

Night or day. The immense choruses come at night and the vast majority
of mating must come at night. Some rainy or showery days or humid over-
cast diurnal weather may make them active in mating. We have no positive
records of mating being initiated by day though when crests come no doubt
some mating carries over into the day or begins about dusk or late afternoon.

Amplexation. {Normal, abnormal). I have seen only two embraces of
this species, the first one pectoral, the second abnormal. No doubt the first
is normal. In the evening of April 29, 192 1, I have this entry: "In Long Pond
I heard R. grylio. The males were out in the middle of the pond among the
Pontederia. The R. grylios can easily be picked up with a flashhght. Some-
times a frog will move a foot or two or three. They rest on the surface. If
one keep quiet and put out the flashlight you can hear them jump once in a
while on the surface. Took a female R. grylio where there were several males.

374 Frogs of the Okefinokee Swamp

Concluded I might mate them. Brought them back and put them in a fish
can (absolutely dark within)." The following morning we noted "This
morning at 6 the Rana grylios were mated." We tried later to photograph
them but they broke the embrace.

One journal entry is our only evidence. On May 21, 192 1, we have
"After supper I went down to Billy Lake Landing. R. grylio going strong.
Air 7o°F. at 7 p.m. Tonight at 8 we went to Chorophilus pond and Engy-
stoma ditches. Air 70°. In the bays around Billy Lake at 12 midnight a
wonderful R. grylio chorus. In the Engystoma ditches caught a female R.
grylio which was in the middle on trash. She jumped towards the vicinity
of a male. Caught her. Later caught a male. Put them together. He mated
axillary. Later with one arm behind and one ahead of forearm. Once his
finger seemed to extend down as in a normal Rana male but this is normally
axillary in this male."

OVULATION

Habitat. (See non-breeding habitat.)

Period. In 19 12 report we note of our 7 specimens only two are adult
females. One was taken on the Suwannee River, June 17, and the other on
Floyd's Island Prairie June 26, 191 2. Both were spent and this would indicate
earlier breeding. It appears that the breeding may come in May and through
June because of the chorus June 18 and the fact that males taken from June
1 5- July 15 or even later had the first finger strongly swollen. At the herony
we took tadpoles 18 mm. long which we took to be this species." In 1923 the
author reported (Copeia, 1923, p. 34) R. grylio as one of five species to begin
breeding from May 15 on to June i. In July- August 1924 (p. 379) Mrs.
Wright and I pronounced the "Season, May 24 to August 21."

Inasmuch as we not uncommonly hear them in chorus or abundance until
September i just before darkness comes on, all through the night and at day-
break we suspect this species to breed until September i. Our latest record
of numbers comes September 5. We actually found eggs on May 24, 192 1,
May 28, 1921, June 8, 1921, June 10, 1921, July 28, 1922.

Temperature and humidity. On May 24, we found our first masses. My
records for that day are somewhat as follows: "In Long Pond this morning
I found where we formerly photographed the male R. grylio, two masses of
R. grylio eggs. They were about ready to hatch. Must have been laid the
night of May 21-22 or that of May 20-21. In another place in middle of
pond found two more masses probably laid night of May 22-23. At 10:30
a.m. temperature in air i foot above water 94°, water's surface 85°. The Cox
boys report having found May 2 1 in afternoon a fresh mass of R. grylio eggs
along the edge of Billy's Lake." This coincides with our belief for the ovula-
tion of the record for May 24. On May 2 1 it rained continually in the swamp;
our temperatures were not below 70°; those outside ranged from 67-80° and
rain was general at St. George, Hilliard, Waycross, etc. The temperatures
for localities outside the swamp for four days preceding May 24 were minima
62-69°, maxima 78-94°, our temperatures 64-94°.

Rana grijlio 375

The film of May 28 probably was laid May 25 when rains were general at
St. George, Hilliard and Waycross. For four days before May 28 minima
ranged from 60-70°; maxima, 87-97°. Our temperatures were 64-94°.

On June 8 the fresh eggs were not laid in rainy weather. The tempera-
tures for 4 preceding days on the outside of the swamp were minima 56-70°,
maxima 84-87°. Our temperatures were 63-82°.

The next record June 10, 192 1, was preceded for four days by maxima
75-90°, by minima 55-70° for outside stations and by our temperatures 63-
84°, and no percipitation for 4 or 5 preceding days.

In 1922 on July 28 found them breeding. For several days before July
28 it had rained. For four days before the nearby Georgia and Florida
stations had minima 72-76°, maxima 91-96°. These general conditions of
72-76° precipitated general choruses.

In general we expect this species to breed when the air is humid, and nights
and general temperature conditions are 63-70°, but the optimum period is
when minima are from 68-78° or higher and maxima mount to 85-96°.

Egg-laying process. No actual observations on the egg-laying process
have been made by us or is in print to my knowledge. The film being similar
to that of the greenfrog or bullfrog is doubtless laid in the same fashion.

EGGS

Attachment. The eggs are usually attached to pickerel weed (Pontederia)
stems in midpond, in the cypress pond on the islands or to "bonnets" on the
prairies or amongst the maiden cane along the water courses.

On May 28, 192 1, we found them amongst pickerel weed and lizard's
tail or in the outer edge of the Saururus Zone or attached to flag (Iris). On
June 10, 192 1, we had to make a detour to the west of Minne Lake (then
filled from lumbering) in a cypress bay. Beside the temporary boat trail in a
rather open place among the cypress we found a mass amongst saw grass,
maiden cane and wampee.

On July 28, 1922, in a cypress pond outside of the swamp near Station
Branch, found a mass hatching near the outer edge of a mass of arrow-leaved
species of plant.

Egg tnass. On June 12, 19 17, at Theodore, Alabama, we were informed
that the large southern bullfrogs which we had captured "laid small beadlike
eggs in large masses on the surface of the water."

The masses found May 24, 192 1 were films at the surface of the water.
These films are generally one or two eggs thick. One mass was 15x18 inches
in diameter; another 12 x 12. All four masses were at least a foot in diameter.
The mass the Cox boys found May 21 was also a film. On May 28, 192 1, the
first mass was 10 to 12 inches x 18 to 20 inches; the second 15 x 10 inches. On
June 10 'Bud Carter brought me in some fresh bullfrog (R. grijlio) eggs. They
found them in maiden cane in Billy's Lake. He said the mass was "15 inches
across it." On July 28, 1922, found a film about 16 x 16 inches in diameter.

In 1920 we wrote (p. 26) "The eggs of the southern bullfrog are not known,
and we are not very familiar with its breeding habits. In Georgia, Florida

376 Frogs of the Okefinokee Swamp

and Alabama the writer has chanced upon this species in full croaking season
in June and July. This species is reputed to lay small eggs in large masses on
or at the surface of the water in the early summer, and if this be true the habits
of this form are closely similar to those of the northern bullfrog."

In 1923 (p. 34) the author asserts "And Rana grijlio has large films, one
by two and a half feet across much as does Rana catesbeiana.^'

In 1924 (p. 379) A. A. Wright and I state the "egg mass (to be) over i
sq. ft. in area (144 to 288 sq. in. or 900 to 1800 sq. cm.) or 12 by 12 inches to
12 inches by 25 inches in diameter; usually in midpond; ..." Some Rana
grylios reach a size (almost as large as Rana catesbeiana and no doubt lay films
of about the same size as occur in the common bullfrog.

Egg description (Field). The first fresh eggs secured June 8 were measured
roughly at night with a lantern light lest we lose the record. These records
were "vitellus 1.6 mm; inner envelope 3.6.; outer envelope 4.5-6 mm. These
will be revised tomorrow." On the morrow, the measurements were made as
follows: "vitellus i 55-1.85 mainly 1.6 mm.; a vitellus of 1.6 has vitelline
membrane 1.8 across it ; outer envelope 6 mm., sometimes 7 mm., some 4 mm. ;
inner envelope 3.6-4.0 mm.; vegetative pole white, animal pole black."

No eggs were laid in camp by captives and our identification is presump-
tive but strongly supported. No R. catesbeiana are present. The only form
which might prove confusing are individual R. clamitans eggs. Whenever we
visited the places where the two were present R. grylio was in the middle of
the pond while R. clamitans was always in the wooded tangles at the edge of
the pond — Rana grylio in open pond, Rana clamitans in the most shaded parts
near the land's edge — distinctive habitats.

In other field notes we have: outer envelope 3.8-4.0 mm.; 4.5-6 mm.;
4-6 or 7 mm.; inner envelope 2.8-3.2 mm.; 3.6 mm.; 3.6-4.0 mm.; vitellus,
1.8 mm.; 1.6 mm.; i. 55-1. 85 mm. After careful laboratory checks the measure-
ments when summarized are:

Outer envelope 3.8-7 mm., merging more or less into the general glutinous
mass; inner envelope 2.8-4.0 mm.; mode 3.0 mm., average 3.45 mm.; vitellus,
i. 4-2.0 mm., mode, 1.8 mm., average 1.7 mm. Egg complement, 8,000 to
15,000.

In the possession of inner envelope the egg of Rana grylio appears more
like that of Rana clamitans than that of Rana catesbeiana.

Dangers. Sometimes one can see a spoiled mass very readily. On May
28, 192 1 "in a pond southwest of the boys' swimming hole found at the surface
of the water amongst pickerel weed (Pontederia) and hzard's tail (Sanrurus)
a mass of R. grylio eggs. I saw it some distance away because of the bluish
appearance of spoiled eggs. Sometimes a mass is revealed by a few eggs dried
or drying above the water on an iris or pickerel weed stem. Occasionally the
cypress ponds raise considerably after eggs are laid and some of the films may
be 3-8 inches below the surface. The most serious condition is where sudden
sinkings of level leave eggs attached far out of the water. Where most of the
eggs are laid, namely, on the prairies the loss probably is not so serious as in
isolated habitats. I know of no animal enemies which feed on the eggs.

Rana grijlio 377

HATCHING PERIOD

The eggs of the morning of May 24 were about hatching on that date. If
they were laid on the night of May 21-22 or on that of May 20-21 we would
have 56 to 78 hours elapsing previous to hatching. On May 21 on Billy's
Lake one of the boys found fresh eggs in the afternoon. Those eggs found May
24 were probably laid at the same time or about 78-80 hours after deposition.
With temperatures of 65-95° obtaining it is easy to expect that 2-3 days
would be a normal period for these surface eggs. Often in shallow water or
surface water at this time the temperatures were 85-90°.

MATURE TADPOLE

Color description from life. Belly is between citron yellow and pale lemon
yellow or lemon yellow with a prominent reticulation of brownish black. On
the sides the lemon yellow spots are surrounded sometimes by pinkish vinace-
ous or orange vinaceous, or coral pink. All the throat region back to the pec-
toral region is a clear black. Across the pectoral region is apple green or
light bice green. On either side of the throat and below the eye and over the
snout and in front of nostril are spots of belly color. Over the back overlaying
black spots and between them is forest green or dark green or elm green.
When a tadpole is young it has a pinkish vinaceous line of spots or stripe on
the upper jaw to be.

Tail. The lemon yellow spots on the sides are also on the base of the
muscular part of the tail as more or less rounded spot with more prominent
pinkish vinaceous. About an inch from the base of the muscular part the
spotting goes along on to the middle of the muscular part for an inch or less
as a long band of pinkish vinaceous or as a row of spots. Below this band are
some rounded pinkish vinaceous spots like the band. Those extend on to the
lower crest. More or less of a long black line of spots above this pinkish
vinaceous band. This black line starts in the middle of the muscular part
but as it approaches the tip of the tail it bends down toward the muscular
black line. LTpper crest with no yellow or whitish spots. Lower crest with
many spots for first two inches.

Pupil yellow or bronzy-rimmed. Iris flecked all over with bronzy. Back-
ground of iris purplish. One looked greenish like the body color.

General appearance. Tadpole quite large (100 mm.) with black gular
area and speckled belly which is so heavily pigmented the intestine does not
show through in preserved specimens. Ventral contour narrower than in R.
catesbeiana and more of the R. clamitans type. Tail elongate, tip sharply
acuminate. Dorsal crest at widest part about equal or slightly less than width
of musculature, and extending on to dorsum slightly ahead of the vertical
through the buds of the hind legs. Spiracle sinistral, just visible from the
dorsal aspect, obliquely directed upwards and backwards, opening elliptical
but little revealed from side. Spiracle opening just touches lateral axis
(muscular axis). Eye on or just above lateral axis.

Anus, dextral, opening at edge of ventral aspect.

378

Frogs of the Okefinokee Sivamp

Muciferous crypts present but rather indistinct, this species easily sepa-
rated from R. damitans in this regard.

Mouth -parts: Teeth 2 '3. Edge of upper labium about equal or slightly
longer than the length of the upper horny beak, and fringed by a continuous
row of teeth. In either corner, beneath this fringe is only one very short row
about one-sixth to one-eighth of the length of the upper fringe. Median
space between these two short rows is two and one-half to four and one-half
times the length of either of these rows. The outer end of the second upper
row does not extend be3^ond the end of the upper fringe of teeth. The third
lower labial row of teeth much shorter than the single separate row of papillae,
much more than length of horny beak (contained i| times in it) and 13
shorter than the second or first lower rows of teeth. These two rows about
equal to horny beak in length and equal to each other. The first row may be
continuous or interrupted in the middle. In mouth parts it is distinctly of
the R. damitans type and nearer this species than R. catesbeiana.

Measurements. Length of body (31.0-35.6 mm.) in tail (55.0-70.5 mm.)
1.7-2.4, average 1.84; width (17-22 mm.) of body in its own length 1.5-2. 15,
average 1.8. Depth (14-20 mm.) of body equal to (i.o) or shghtly less (.S-.g)
than body width. Depth of body 1.5-2.55 in body length, average 1.98.
Depth (17-23 mm.) of tail in length of tail, i. 45-1. 8, average 1.7. Depth of
tail 1. 05-1. 5 deeper than depth of body, average 1.2. Muscular part (9.5-
14.5 mm.) contained in depth of tail 1.3- 1.7, average 1.5; in smaller specimens
1.75-2. 1. Spiracle 1.1-1.6 nearer base of hind legs or vent region (13-19 mm.)
than the tip of the snout. Spiracle 1.2 5-1. 9 nearer eye than base of hind legs
or vent, average 1.56. Eye 1-1.35 nearer to tip of snout (7-10 mm.) than to
spiracle (8-12.5 mm.); in some younger tadpoles occasionally near (.88-95)
spiracle. Nostril i. 0-1.4 nearer snout than eye, rarely equidistant. Mouth
(4-6 mm.) usually 1.-1.6 larger than internasal space (3.6-5.5 mm.), average
1. 15. Mouth contained 1.5-2.37 (average 1.94) in interorbital distance (7-
12.0 mm.). Internasal space contained in interorbital space 1.8-2.6, aver-
age 2.16.

The dimensions of the largest tadpole are:

mm. mm.

Total length
Body length
Body depth
Body width
Tail length
Tail depth
Musculature of tail
Spiracle to snout

100. o Spiracle to vent 19

36.6 Spiracle to eye 10

20. Eye to snout 9

22.0 Eye to nostril 5

64 . 4 Nostril to snout 3
23.0 Mouth 5

13.5 Interorbital distance 1 1
21.5 Internasal distance 5

General remarks. The large tadpoles are difficult of capture. Usually it
is impossible to haul a minnow seine where they occur. Occasionally when a
cypress pond dries up, they can be caught in the centrally cleared space, the
normal alligator hole of the pond. Rarely one catches them in the scoops of

Ranagrylio 379

a dip net and smaller stages frequently come in the floating maiden cane
masses we often pull over the gunwale into the boat. Rarely one chances on
shallow water areas cut off from cypress ponds, and in these localities we had
our best luck. We found few mature tadpoles.

LARVAL PERIOD

On August 3 and 4, we found in two different ponds "plenty of R. grylio
tadpoles of two different sizes. Does it stay two years as a tadpole or only
one year as in R. clamitans? Some of them seem still to retain a black rim
on the lower crest ridge." Our largest R. grylio tadpole reaches 100 mm. and
transformation ranges from 43-51 mm. Rana catesbeiana tadpoles may reach
140 mm. and transform at 43-59 mm. Rana clamitans tadpoles may reach 84
mm. and transform at 28-38 mm. R. septentrionalis tadpoles may reach
99 mm. and transform at 29-40 mm. In the north R. catesbeiana takes two
years for transformation but some Rana catesbeiana sent by Viosca to Japan
developed in one year of tadpole life. Do those of Louisiana do the same?
If so possibly Rana grylio may transform after one year of tadpole life. It
may take two years in some instances even in Gulf Coast environment.
Some of the Rana clamitans 70-84 mm. tadpoles winter over to transform at
28-38 mm. Then possibly to attain greater growth R. grylio may take a
slightly longer period. In the South generally development progresses faster
and breeding periods are longer. For example, in the Northern States we
would be surprised to have R. catesbeiana breed in February as it did in San
Antonio parks February 1925.

TRANSFORMATION

Period. In 191 2 we had "only one transformed specimen which measures
44 mm. It was taken May 30, 19 12, from the stomach of a snake."

In 192 1 we found a transformed frog April 24.

On April 25, 192 1, we caught a young recently transformed R. grylio
amongst the Pontederia. Later caught a second with a stub of a tail present.

In a cypress pond which had almost dried up we found transformed southern
bullfrogs. In the center was a small circular water area. "Around this small
pool was a large moist mud flat where a pig was rooting up frogs, etc. Under-
neath the mats or carpets of grass which originally was in the water found
little transformed R. grylio in small moist pockets dug in the mud. With them
were Manculus quadridigitatus, a salamander, a ring-neck snake (Diadophis
punctatus) and a skink. In the pool were pygmy sunfishes (Elassotna) tad-
poles of Rana sphenocephala and water beetles. Florida Blue Herons and
Florida Crackles as well as the pigs were having a feast at this pond." No
doubt the young southern bullfrogs suffered.

On May 6, 192 1, Dave Lee "caught three R. grylios of three sizes from
Billy's Lake: i transformed; one intermediate; and one adult male." On
May 9 two more were taken; on May 10 we saw plenty of small southern bull-
frogs in one pond. On June i we saw several just transformed southern
bullfrogs in a pond outside the swamp. On June 15 we had some tadpoles

380 Frogs of the Okefinokee Swamp

transform. On June 19 we have a record of one transformation. Beside
the above notes from our Journal we have the following dates for transforma-
tion: May I, June 20, July 11, July 19.

In 192 1 our dates extend from April 2 4- July 19.

In 1922 on June 14 when we arrived at Ghesser Island, there were several
transformed frogs found. In the same place two weeks later we still found
transformation in process.

Size. In 1920 we held "Some of the southern bullfrog tadpoles may
transform in the very last of May and the minimum size recorded is i 3/4
inches." The transformation size of our lone specimen of 19 12 measured 44
mm. At the same time May 30 we secured another specimen 53 mm. in
length of body. In 192 1 we have the following measurements: On April 24,
we have two transformed at 43 mm. (C. U. 266) and 51 mm. (C. U. 265).
On April 25, two were captured and measured 48 and 47 mm. respectively.
Another of April 25 measured 53 mm. but it is, however, past transformation.
On April 26 one (C. U. 213) measures 43 mm. One was captured May 6 but
is not at hand. Another (C. U. 291) found June 19 is 45 mm.

Besides the above notes from our Journal we have the following specimens:
May I, two at 37 and 39 mm. respectively; June 15, one at 32 mm. trans-
forming but progress not completed; July 11, two at 38 and 47 mm. respec-
tively; and July 19 two at 41 and 43 mm. respectively.

In 1922 we secured transformed individuals as follows: June 14, one at 49
mm., July 17 one at 42 mm.

In general in 191 2 we have a 44 mm. specimen; in 192 1 specimens from
32-48 mm.; in 1922 specimens from 42-49 mm. The measurements of 32-49
mm. about represents the size at transformation, although we believe the 32
mm. rather small for a normal R. grylio transformation. This places this
species intermediate between the transformation size (28-38 mm., average
(Ithaca) 3.2 mm.) size of Rana damitans and the size (43-59 mm., average
53 mm. (Ithaca) of Rana catesbeiana.

General remarks. When the hind legs are well formed the upper fringe is
generally gone and the upper lateral ridge of teeth also, but some of the lower
labial teeth may yet be retained.

These young frogs after transformation stay in the same habitat as the
adults. We captured them by hand at night with flashlight amongst the
bonnets of cypress ponds. Occasionally they could be caught along the maiden
cane edges with a dip net and torch light. On July 7, 1922, we found many
of the young ones in the shade of the small wooded islets (houses) of the
prairies. They would start off with a startled cry as one approached and in
this respect reminded one of young Rana catesbeiana.

On May 14, 192 1 , at 8 p.m. we found plenty of the young transformed frogs
and somewhat larger young frogs resting on the sphagnum mat or vegetation
of a cypress pond and caught several of them by hand.

On July 17, 192 1, on Chesser Island we took a 40 mm. Rana grylio. Two
days later in the same place we caught a mature male R. virgatipes (vocal sacs
and thumb developed) of 43 mm. They looked very much alike but the fourth

Rana gnjlio 381

toe character separates them. The male R. virgatipes has a more pointed
head, much thicker legs, broader body and heavier darker ventral spots with
larger white interspaces. The dark speckled or spotted venters are typical of
transformation size in R. grylio, R. catesbeiana, R. clamitans and R. virgatipes
and as growth begins all come to have venters with larger dark spots with
larger light interspaces. In R. virgatipes the 2nd finger is equal to or less than
the first finger, in R. grylio the second finger is usually larger than the first
finger.

GROWTH

Of the 191 2 collection we wrote as follows: We have only one transformed
specimen which measures 44 mm. It was taken May 30, 19 12, from the
stomach of a snake. A specimen taken May 30, 191 2, on Honey Island
measures 53 mm. and doubtless represents a year old, while another taken
at the same time and measuring 72 mm. looks to be a two year old. Another
taken June 15, 1912 on Billy's Island measures 88 mm. or one from Floyd's
Island June 26, 1912, measures 88 mm. These doubtless are three year olds.
Three others taken in the same summer measure 96. 5-1 01 and easily are four
years old. These measurements 44, 53, 72, 88, 96.5-101 fairly well represent
the growth for 4 years after transformation.

Some of our records for 192 1 follow: On April 24 and 25 we secured seven
specimens 100, 92, 95, 82, 50, 44, 44 mm. respectively, the last two at 44 mm.,
being transformed individuals. They seem to fall into 4 groups 100, 92-95,
82, 44-50. From April 22-May, 6 sizes: one female 112 mm.; two males of
99 and 100 mm. respectively; two males 94-97 mm. respectively; one female
83 mm.; another 77 mm.; and two transforming at 37 and 39 mm. Those
taken May 10-23 f^l^ iiito three groups: one at 91 mm., the next group with
86, 77, 76 mm. respectively; the last group with 58, 62 mm. respectively.
June 15-20, 192 1, we have transformed example 46 mm. and one 73 mm.
Specimens taken July 11-19 are 38, 41, 43, 47, 61, 67, 67 mm. respectively
or of two groups.

In 1922 from June 14-August 9 we took specimens 42, 49, 56, 60, 61, 69,
80, 84, 84 mm. in length.

The 1921-1922 material seems to fall into the following groups:
(i) 32-49 mm., average 41 mm. tranformation.

(2) 56-69 mm., " 62 . 5 mm. one-year-olds.

(3) 73~86 mm., " 80 mm. two-year-olds.

(4) 91-102 mm., " 96 mm. three-year-olds.

(5) 107-115 mm., " 1 1 2 mm. four-year-olds.

This species may reach 150 mm. Dr. Stejneger's type (U. S. N. M. No.
27443 is 125 mm. and U. S. N. M. No. 30951 is 148 mm.).

Since this paragraph was written we have examined some specimens Dr.
Barbour has. Some of these are as follows 120 mm. cf, 125 mm. 9, 136
mm. d^, 143 mm. 9 , 161 mm. 9 and 161 mm. 9 . These are bigger than any
others in collections. The author, however, has also seen equally large indi-
viduals in the Okefinokee Swamp.

382 Frogs of (he Okefinokee Swamp

FOOD
We have not examined the sixty of more individuals we have for food. I
know of no records in print.

ENEMIES
On the basis of our 191 2 specimens we wrote: Out on the open prairies
the southern water snake {Natrix fasciatus fasciatus) is a bitter foe of the
southern bullfrog. One water snake had taken nearly two transformed frogs
of this species. Along the bonnet "runs" the moccasin and pied water snake
make life miserable for these frogs. Among the birds, the herons, and ibises
relish them. We have not examined our snakes, turtles or fish of the 192 1
and 1922 collections and are unable to determine which fed on southern
bullfrogs.

AUTUMNAL DISAPPEARANCE

If it be like a bullfrog in the South, it will spend four or five months in
hibernation. The months from November i -March i or later are doubtless
usually inactive for the species. In some cases they go into hibernation be-
fore November i or come out before or after March i as does Rana catesheiana.
For example, I have heard Rana catesheiana croaking in the first or second
week of February in Texas.

AFFINITIES

Indirectly Dr. Stejneger indicates the affinities of Rana grylio with Rana
catesheiana by the title of his paper "A New Species of Bullfrog from Florida
and the Gulf Coast." In diagnosis he pronounces it "Similar to Rana cates-
heiana, but with the fourth toe much shorter in proportion; the third toe,
measured from the inner metatarsal tubercle, being more than three and one-
half times the difference between the third and fourth toes." Further in his
description he says:

"These differences in the relative length of the toes, being capable of the
most concise definition, have been utilized primarily for the characterization
of the new species, but there are numerous other features which prove it to
be very distinct from R. catesheiana. Thus, for instance, the snout is much
shorter and less high, so that the nostrils appear to be nearer the tip of the
mouth; the head is also narrower behind; the vomerine teeth are very close
together, with hardly any space between two patches; the tongue is much
less broader and thinner, with remarkably long and thin 'horns,' which are
very far apart; the color is apparently much darker brown, though there may
be R. catesheiana nearly as dark. Add to this the difference in voice and we
have clearly one of the most distinct species of frog in the United States."

"The general habitus of R. grylio is that of Rana catesheiana, including the
large tympanum and the absence of a dorso-lateral glandular fold. It is also
a large frog, though whether it reaches the extreme size of R. catesheiana may
be doubted."

In 19 14 R. F. Deckert holds it to be "a beautiful frog closely related to the
common bullfrog," while Miss Dickerson calls it a frog "really very different
in appearance from the common bullfrog, not resembling it in shape, proper-

Plate XXXVIII
River-swamp frog (Rana heckschen)

1 . Thompson's Landing, St. Mary's River, Ga. July 9, 1922.

2. 3. Mature tadpoles, Alligator swamp, Callahan, Fla. August 18, 1922. x i.e.

4. Mature frog, small size, 56 mm. Alligator swamp, Callahan, Fla. August 17,

1922. Lateral aspect, x 0.75.

5. Mature frog, large size, 95 mm. Alligator swamp, Callahan, Fla. August 17,

1922. Lateral aspect, x 0.66.

6. Mature frog, large size, 95 mm. Alligator swamp, Callahan, Fla. August 17,

1922. Ventral aspect, x 0.5.

Rana grijlio 383

tions of body or coloring. The long narrow, pointed head, with its large eyes
set close together, tell the story at once. But other evidences lie in the ratio
of the lengths of head and body (...), in the greater length of toes (except the
fourth) (. . .), and in the finely pitted texture of the skin. In addition to all
these points, there is that of coloration. . . . Rana grylio shows close relation-
ship to Rana virgatipes of New Jersey in shape and proportion, metallic
coloring, texture of skin, large size of eyes and ears, in lacking the lateral folds,
and in aquatic habits."

In 19 14 the author in the light of his 19 12 experiences with Rana grylio
made the following unfinished memoranda: "In many ways Ratia grylio ap-
proaches Rana virgatipes. Like the latter it has on the posterior faces of the
hind limbs alternating horizontal bands of dark and light colors. The general
mottlings of the venters and the coloration of the hind limbs to toe tips are
almost duplicate. The webbing of the hind toes is more or less similarly
mottled. The fore limbs of Rana grylio are more or less unicolor without the
characteristic dark anterior faces of R. virgatipes. Both have no dorsolateral
glandular folds. The four dorsal stripes of R. virgatipes are missing in R.
grylio and R. virgatipes has two phalanges of the fourth toe free." In 1924
(p. 141) in discussing Rana heckscheri we held "These southern bullfrogs or
' Joe Browns' are very distinct in adult, tadpole and egg characters from the
bullfrog, Rana caiesbeiana."

In 19 16 Dr. T. Barbour writes: "I think our famiharity with Rana grylio,
the big frog found swimming among the 'bonnets' in some of the Florida
lakes, has made this frog's relationship to the New Jersey sphagnum frog
seem doubly striking, for their coloration in life is astonishingly similar, and
in fact one seems to be but a miniature representative of the other."

In 1920 Dr. Boulenger (p. 425) thought Rana catesbeiana and Rana grylio
near the intermediate form from which the other Ranas of North American
come rather than from Rana septentrionalis (Cope's belief).

Rana grylio adults surely, as Dr. Barbour states, seem big editions of Rana
virgatipes in general coloration and general habitus. A young transformed
Rana grylio might more quickly be confused with an adult or half-grown Rana
virgatipes than with any other species of Rana. Their ranges overlap in the
Okefinokee. Both are very aquatic species and Rana virgatipes might be held
the northern offshoot. Usually we say R. grylio has not the two or four lateral
yellow brown or golden streaks on the body as has Rajia virgatipes. But I
have seen several recently transformed R. grylios with light streaks from eye
backward where a dorsolateral fold would be if present. And Viosca (1918,
p. 161) writes "Four hghter longitudinal bands of orange brown are sometimes
discernible on the back, especially in young specimens."

In egg mass it has a film like Rana catesbeiana and R. claniitans, but in egg
characters it has an inner envelope of jelly like Rana clamitans. In tadpole
the body seems more like Rana clamitans but in size it is midway between
R. clamitans and R. catesbeiana. In mouth parts it is more of the R. clamitans
type. In transformation it occupies a median position between the two species.
It belongs in the Rana clamitans-catesbeiana group with close affinities also for
R. virgatipes.

384 Frogs of the Okefinokee Swamp

BIBLIOGRAPHY

1916 Barbour, T. Copeia, Jan. 24, 1916, No. 26, p. 6.

1920 Barbour, T. Copeia, July 31, 1920, No. 84, p. 55.

1 791 Bartram, William. Travels through North and South Carolina, Georgia, East and
West Florida, etc. Phila., pp. 276, 2jj.

1920 Boulenger, G. A. Am. Acad. Arts Sci., Vol. 55, No. 9, August, 1920, pp. 415, 4.21, 422.
1914 Deckert, R. F. Copeia, Feb. 14, 1914, No. 3.

1914 . Copeia, April 15, 1914, No. 5.

192 1 . Copeia, Mar. 15, 192 1, No. 92, p. 20.

1922 . Copeia, Nov. 20, 1922, No. 112, p. 88.

1906 Dickerson, M. C. The Frog Book, N. Y., pp. 226-227. Plates LXXXV, LXXXVI.

1922 Lading, H. P. Geo. Survey Ala Mus. Paper No. 5, p. 20.

1923 Pratt, H. S. 1923, p. 183.

1901 Stejneger, L. Proc. U, S. Nat. Mus. XXIV, 1901, pp. 21 1-2 15.

1918 Viosca, Percy, Jr. 3rd Biennial Rept. La. Dept. Conserv. pp. 161, 162.

1923 . Copeia, Feb. i, 1923, No. 115, pp. 39, 43.

1920 Wright, A. H. U. S. Bur. of Fisheries Rept., 1919, Doc. No. 888, pp. 10, 19, 21, 26, 34.

1923 Wright, A. H. Copeia, Feb. i. No. 115, p. 34.

1924 . Proc. Biol. Soc. Wash. Vol. 37, pp. 141-152.

1923 Wright, A. H. and A. A. Wright. The Anatomical Record Vol. 24, No. 6, Jan. 20.
1923, p. 406.

1924 . Am. Naturalist, Vol. LVIII, July-August, 1924, pp. 376, 377, 379.

Rana heckscheri Wright

(PI. Ill, Fig. 4; XI, Figs. 7, 8; XIV, Figs, i, 2; XVII; XXXVIII)

COMMON NAMES
Greenback. Heckscher's Frog. River Swamp Frog

RANGE

Check list. Described since Stejneger & Barbour's 2nd edition.

Supplementary records. The type locality where first recognized (1922)
is Alligator Swamp, Callahan, Florida. The known range is from Savannah,
Georgia to Fargo (on Suwannee River) and Folkston (near St. Mary's River)
in Georgia, all along the St. Mary's River on both Georgia and Florida sides
and in Florida to Callahan. In 1928, June 9 and 10, we visited the type
locality and also found them in Mr. Davis's clay pit. It ought to be farther
north along the coastal strip to Wilmington and westward to Louisiana.

In American Museum of Natural History we found three specimens taken
by T. Hallinan. One, a male (No. 16096), 95 mm. long, was taken at East-
port, Florida and two more, males (Nos. 16666-67) 97 and 91 mm. long were
secured near Jacksonville, Florida.

In a recent letter (Nov., 1929) Mr. O. C. Van Hyning writes me that he has
found 12 miles from Gainesville, Florida, at Waldo, some tadpoles of Rana
heckscheri. His captures he compared with some freshly caught tadpoles of
Rana heckscheri which I left with him at Gainesville, Florida in June 1928.
In June 1928 we captured many intermediate sizes of this species but these
like some of the 1922 specimens are now missing.

Since the three preceding paragraphs have been written two more records
have come in recent letters. On August 19, 193 1, Mr. O. C. Hyning who
had formerly found this species in Georgia west of Okefinokee Swamp in
1930, writes that "he has a Rana heckscheri adult from 2 miles east of Gaines-

Rona heckscheri 385

ville, Florida." On August 15, 193 1, Morrow J. Allen, of the Caribbean
Biological Laboratories, Biloxi, Mississippi, sends me "under separate cover
two specimens of a tadpole that were taken last month in the overflow of a
stream along with about 25 others. I have never seen this tadpole before
and ask if you will give me the identification. I notice that the plate of the
tadpole of Rana heckscheri in your synopsis of North American tadpoles some-
what resembles it." They are Rana heckscheri tadpoles.

Local Okefinokee records. No adult specimens have been taken within the
swamp.

GENERAL APPEARANCE

It looks like a big Rana clamitans with melanistic and sulphur venter. It
has more rough dorsum than Rana clamitans, Rana catesbeiana or Rana grylio.

COLORATION OF SPIRIT SPECIMENS (1922)
In spirits, four 95 mm. males of four species are as follows: bister or mummy
brown on dorsum of R. catesbeiana, brownish olive in R. grylio, deep grayish
olive in R. clamitans, and deep mouse gray in R. heckscheri; upper parts with-
out very distinct dark spots in R. clamitans and R. cateheiana, with prominent
large black spots in R. grylio, and with many small dark spots R. heckscheri;
venter of R. clamitans clear white except for the yellow throat, venter of R.
catesbeiana heavily blotched with black, so also in R. grylio — all three, how-
ever, with a white background color but in R. heckscheri the deep mouse gray
or dark color so prominent it becomes the background color and the white,
scattering spots; light spots on upper and lower jaws more prominent than in
R. clamitans. Throat in males prominently deep mouse gray, deep neutral
gray or violet slate while female's throat is whitish.

MEASUREMENTS

(Recent Material)

Head to angle of mouth 1.2 (36 mm.) — 1.25 (44 mm.) — 1.2 (56 mm.) —
1. 31 (82 mm. cf) — 1. 17 (95 mm. cf") — 1.2 (113 mm. cf — 1.16 (125 mm. 9)
in width of head; head to rear of tympanum i. 0-.93-1. 04-1. ii-i. 08-1. 09-1. 09
in width of head; head to angle of mouth 3.0-3.-3.66-2.8-2.82-2.7 1-2.82-3.0
in length of body; head to rear of tympanum 2.5-2.75-2.43-2.41-2.5-2.56-2.9
in length of body; snout .84-1.0-.9-.71-.69-1. 0-1.05 in fourth finger; snout
1. 0-1.35-.8-.78-.81-1. 06-1.05 in first toe; eye i. 4-1. 07-1. 43-1. 55-1. 6-1. 36-
1.4 in snout; eye .60- 54-. 70-1. 3-1. 3-1. 3 6-.85 in tympanum; eye i. 2-1. 08-
i-3~i-33~i-3~i-36-i.4 in first finger; tympanum 3.66-3.86-3.1-1.5-1.61-.53-
2.2 in intertympanic width; tympanum 2.33-2.0-2.0-1. 16-1. 23-1. 0-1.65 in
snout; internasal width 1.0-.9-1. 08-1. 43-1. 07-1. o-i.o in upper eyelid width;
interorbital width 1.0-1.0-2.0-1.14-.83-1.36-1.2 in upper eyelid width;
interorbital width i.o-i.i 1-1.83-1.0-.77-1.36-1.2 in internasal width; inter-
orbital width 2.75 3. 0-5. 16-2. 57-2. 33-1. 3-3. 25 in intertympanic width.

Forelimb: 2.0-2.0-1.93-2.34-2.0-2.5-2.2 in length (snout to vent) ; forehmb
2.72-2.9-2.9-3.4-2.83-3.5-2.8 in hind limb; first finger i. 33-1. 43-1. ii-i. 08

386 Frogs of the Okefinokee Swamp

1. 23-1. 23-1. 3 in third finger; second finger i.o-i.o-i. 375-1. 18-1. 09-1. 06-1. 05
in first finger; third finger 1.125-1.3-1.2-1.36-1.3-1.3-1.16 in second toe;
fourth finger 1. 1 6-1. 3 5-1. 1 4-1. i-i.i 8-1. 06-1.0 in first toe; fourth finger 1.33-
1. 43-1. 42-1. i-i. 45-1. 23-1. 2 in third finger ;internasal width 1. 5-1. 4-1. 63-1. 7 1-
1. 85-1. 8-2.0 in first finger; i. 5-1. 4-1. 36-1. 57-1. 71-1. 7-1. 9 in second finger;
2. 0-2.0-1. 8-1. 85-2. 3-2. 2-2. 5 in third finger; i. 5-1. 4-1. 27-1. 43-1. 57-1. 8-2.1
in fourth finger.

Hindlimb: length i. 36-1. 46-1. 5-1. 45-1. 4-1. 4-1. 28 in hind Hmb; tibia
2. 5-1. 9-1. 93-2.0 2. 11-2. 0-2. 17 in length; tibia 2.8-2.7-2.89-2.9-2.95-2.86-
2.7 in hind limb; tibia 1.03-.95-1.0-.853-1.04-.80-.96 in forelimb; tibiai.03-
i.o-i. 03-1. 04-1. 06-.93-.95 in hind foot; first toe i. 26-1. 45-1. 5-1. 63-1. 61-
1.5-1.4 in second toe; i. 85-1. 84-1. 87-2. 72-1. 84-2. 1-2. i in third toe; 2.57-
2.2-3.37-3.45—3.23-2.9-2.7 in fourth toe; i. 8-1. 7-2. 25-2. 63-2. 3-2. 1-2.0 in
fifth toe; second toe 1.44-1.27-1.41-1.61-1.14-1.4-1.5 in third toe; 2.0-1.6-
2.25-2.11-2.0-1.9-1.9 in fourth toe; i. 4-1. 23-1. 5-1. 55-1. 42-1. 4-1. 4 in fifth
toe; third toe i. 38-1. 2-1. 59-1. 31-1. 75-1. 4-1. 2 in fourth toe; .96-.91-1.06-.96-
1.25-1.0-.95 in fifth toe; fourth toe 1.0-1.1-1.11-1.13-1.14-1. 0-1.02 in hind
foot; .97-1.1-1. 07-1. 07-1. 07-1. 07-1. 07 in tibia; 1.0-1.05-1.07-.92-1.12-.86-
1.04 in forelimb; fifth toe i. 44-1. 31-1. 5-1. 35-1. 4-1. 37-1. 35 in fourth toe;
internasal width i. 75-1. 9-1. 45-1. 57-1. 85-1. 9-2.1 in first toe; 2.25-2.6-2.18-
2.57-3-0-2.85-3.0 in second toe; 3.25-3.5-3.08-4.14-3.43-4.0 4.4 in third toe;
4.5-4.2-4.9-5.43-6.0-5.47-5.7 in fourth toe; 3.12-3.2-3.2-4.0-4.3-4.0-4.2
on fifth toe.

HABITAT

It seems to be a frog of the swampy edges of rivers and streams, a truly
fluviatile species.

On August 18, 1922, with a light we captured eight or ten frogs of various
sizes from probably one year frogs to full sized adults. We found them in
shrubbery and on the banks about the bases of trees. More were captured
than lost. They were rather awkward in their escape and would tumble off
from their perches. Later in the evening we lost all but three of our capture.

If this were truly a wooded river swamp species, it is hard to explain its
presence in the brickyard clay hole in Callahan. It is only a short block from
Alligator Swamp, the type locality. Either they were introduced or they in-
vaded the place from the nearby Alligator Swamp. The clay hole is immense
in breadth, artificial, yet it is becoming already very swampy in character.
These pools have waterliHes, arrowhead (Sagittaria), water hyacinth, water
pennywort ( Hydrocotyle) and countless other water plants. Some of the
residents told us that when ditches for the clay hole were dug they were
absolutely crowded with seething masses of black tadpoles which must be the
larvae of this species. No other tadpole is so distinctive in the eastern U.S.A.

Cutside and inside the swamp. On the western edge of the swamp the day
we entered the swamp in 191 2 (last of May) "We secured a frog which puzzled
us. We saw it only for a few moments. We soon lost it in the rigors of the
trip." It was a fine male of Rana heckscheri. We have no positive specimen
from within the swamp.

Rana heckschen' 387

GENERAL HABITS

Two journal notes of 1928 (June 8) reveal some of its habits and its habitat.

"June 8 . . . This afternoon went to type locality of Rana heckscheri. . . .
Went along in pickerel weed {Pontederia) on the east side of the bridge (Dixie
Highway) where it is shady. 5 p.m. Finally saw a large frog. Slowly ap-
proached from water side with net in right hand and came close enough to
catch the frog with the left hand. Started skirting several edges. Soon saw
another on the bank amongst pickerel weeds. Crawled towards it on my
hands and knees and caught it. It squealed as do some greenfrogs, bullfrogs
or southern bullfrogs when caught. The children went along and startled
another which gave a startled note of the Rana catesbeiana type. Another
big frog amongst pickerel weed on land and not at water's edge. It leaped
along and into the water before I could capture it. Some three or four lost,
partially because six children trailed along in their anxiety to help. . . . Mrs.
Wright thinks it a different frog. These two frogs we captured are larger
than the three we secured in 1922. They are bigger than any greenfrog. They
have no costal folds. Saw no intermediate sizes."

"June 8. . . Went out tonight to the type locality of Rana heckschen.
Saw a female near the edge of Pontederia bank. Later heard a big frog jump
in. When I approached that area, found a male, a arge one on the bank,
amongst vegetation and not at water's edge. Saw and took another female.
Later in going down a little run where we frightened a Rana heckschen,
heard a male Rana clamitans. Then the boys called me for a large frog they
saw. It proved a female Rana grylio. Went back after the male R. clamitans.
Just before I reached it, in the water resting on a submerged log was a fine
male Rana heckschen. In the shade of trees on the bank was the croaking
male green frog. Small for Rana clamitaris but it croaks. It is different from
Rana heckschen, so also is the Rana grylio female I caught here. This Rana
heckscheri is not Rana catesbeiana though we have not caught the latter here."

This species is not so dextrous as some of its related species. Some of the
boys reported "lots of them. We catch them by hook and with sticks."

VOICE

The discovery of the adults came in this fashion. On August 18, 1922, we
visited Alligator Swamp at Callahan, Florida, because of some curious large
tadpoles we saw there one month earlier. "Mrs. Wright discovered a queer
looking green frog as she supposed and as she was calling to us we were startled
by a call unlike any other Rana we ever had heard. To one it was a snore, to
another a snort, and to others neither. The queer green frog and the author
of the call proved of the same species and not green frogs of which we captured
some for comparison."

Mr. F. Harper, while traveling down the St. Mary's River in August, 1921,
"heard a number of times, in addition to the regular snoring call, a peculiar
explosive snarhng grunt." Then he associated the call with Rana aesopm
but now he believes it this species. He heard it from August 10 or i i-i 7 early

388 Frogs of the OkefinoJcee Siramp

in morning, in mid forenoon, at noon, and in late afternoon, or from about
Moniac to Camp Pinckney.

In 1928 on the afternoon of June 8, we did not hear the regular call of this
species in the type locality. Only the squeal of a captured frog and the
startled note of an escaping frog did we record. In the evening of June 9 "in
the brick yard pools heard no end of Rana grylio, two deep notes. Once in a
while we would hear the snoring notes (3 or 4 of them) of Rana heckscheri."
Certainly the calls were sporadic and this exact period was not a breeding
season for the species.

MATING

Male (From life, August 18, 1922). General dorsal color citrine drab to
grayish olive becoming on top and sides of head and center of tympanum
dark olive buff, Isabella color or cinnamon brown. Ear drum except middle
mummy brown. On back of body and on head and on some of the sides is
some serpentine green. Under parts spotted white with glaucous gray or
light Payne's gray or pale drab on throat and breast. Throat with a little
citron green or deep chrysolite green. Spots on lower jaw rim four or five, sea-
foam yellow, to deep colonial buff in the spot just back of the angle of the
mouth. This spot except above surrounded by black.

Black spot just below angle of mouth to and across the insertion of the
brachium. Three black spots on the front edge of the antebrachium. Rear
of fore legs black to tips of fingers and webs. Tops of the fingers with seafoam
yellow or deep colonial buff spots.

Narrow black bars across the dorsum of the femur, tibia and hind foot.
Rear of femur with white unconnected spots on the bone brown ground color.
Rear edge of hind foot to tip of fourth and fifth toes black.

Iris outside rim bright green-yellow, inner rim capucine orange; and in-
terval black with orange rufous spots.

A younger specimen 65 mm. long is dark olive or deep olive on entire upper
parts and very warty. Throat is deep grayish olive.

Two green frogs from the same place were: one, a female, dark olive buff
uniform; and a smaller one wood brown to avellaneous uniform; back of eye
and side of head and over angle of mouth apple green. Bars on hind legs
absent in female, scanty in smaller specimens. Both with costal folds.

Male {From life, June 10, 1928). Back, top of head, top of fore legs and
hind legs citrine or buffy citrine. Under parts white spotted with olivaceous
black (3) or in lights becoming pale olive gray. So also on under sides of fore
arms and hind legs. Rear of femur with spotting of white or pale vinaceous
fawn. Not in lines of white as in Rana grylio.

Throat with wash of oil yellow or sulphine yellow on a light mineral gray
background. Edge of lower jaw with light spots of white, pale chalcedony
yellow or cartridge buff. Back of angle of mouth and below tympanum is a
spot of light greenish yellow or pale chalcedony yellow. Another one back of
the first. Beginning at angle of mouth an oblique area of black or light
vinaceous drab extends to arm insertion, around it into axilla. A branch goes .
up back of tympanum. Rear of forearm has area of same on dorsum. Webs

Rana heckscheri 389

cloudy with black or olivaceous black (3) or light vinaceous drab. Lower
sides mottled with black and light vinaceous fawn. Sides of face buffy brown.
Tympanum olive brown or natal brown. Center circle buffy brown. A dark
area in inguinal area.

Iris black dotted with orange rufous. Pupil rim cadmium yellow. Iris en-
circled by pale green-yellow or bright green-yellow.

When first caught very black and white beneath.

Another male is olive or dark greenish olive above.

Female (From life, June 10, 1928). Much the same as male but upper
parts much more spotted with black. Head spotted. Tympanum with
blotches of black or olivaceous black (3). Half bars on femur. Bar across
arm insertion. Throat with no or little oil yellow or sulphine yellow. White
spots or pale vinaceous fawn spots on rim of lower jaws prominent. Under-
parts not quite so spotted. Iris same as in male.

Young transformed frogs. (June 10, 1928). A young transformed frog
has a prominent eye. Sometimes brick reddish. (3ften mars orange pre-
dominates over black and iris rim is orange rufous or brighter. Entire under
parts finely speckled white and deep olive gray or deep grayish olive.

Structural differences. The male has very enlarged tympana, enlarged
thumb and doubtless is much more intense in ventral coloration if the younger
underdeveloped males be a criterion or index of the species type. Females
and young undeveloped males more often agree in coloration. Most of these
we lost and can remember them only as lighter in appearance. Probably the
species is of the R. clamitans or R. catesheiana type in amplexation. (1922).

The characters given in 1923 hold but in addition we discover that the
males average somewhat smaller and rougher on the back and sides. Our
eight adults (two of 1922 and six of 1928) are five males and three females.
The males are 82, 95, 102, no and 113 mm. in body length and the females
are 122, 123, and 125 mm. respectively.

Ainplexation. We have no observation. The enlarged thumbs and all
general assumptions or presumptions in the case point to a normal Rana
embrace.

OVULATION

Habitat. See General Habits and particularly General remarks under
Mature Tadpole.

Period. In 1922 we had no positive record of ovulation. Recalling the
small tadpoles of June 16, 19 12, the tadpoles of Aug. 7-16, 192 1, and the small
tadpoles of St. Mary's River July 17, 1922 we then thought the period might
be from June i to July i or 5.

The previous paragraph of 1923 is now supplemented by the impressions
of 1928 (June 8-10). There were transforming tadpoles, 80 or 85 mm.- 100
mm., and some with leg buds slight or none from 80-70 mm. at the type
locality. In the brickyard were transforming tadpoles and the small gold-
banded tadpoles of three groups (10-15 mm., 20-25 nim., 30-35 mm.). These
last three sizes indicate three breedings in the spring previous to June i.
Possibly they may be coincident with previous rainy periods in 1928 if there

390

Frogs of the Okefinokee Swamp

were such and the period of ovulation extends from the last of April or May i
to July 5 or 8. The voice records of June 8-10, 1928, Aug. 10-17, 192 1, and
Aug. 18, 1922 might indicate breeding from June 8-Aug. 18. At none of these
periods were choruses heard. Of the three females taken June 8-10 one was
unspent and two spent. June 10 therefore is not past the ovulation period.
No doubt as in Rana clamitans, R. catesbeiana and Rana grylio this species
occasionally breeds in August.

EGGS

In 1922 we found on July 17 at Thompson landing, St. Mary's River
many small R. heckscheri tadpoles. Then we thought them H. cinerea. It
was not until a month later in another place we found large tadpoles and
adult frogs which evidence revealed that it was a new form, R. heckscheri.
Not until recently did we recall that in a nearby pool (20-30 ft.) away from
the place of the Uttle Rana heckscheri tadpoles did we find (July 9) a large
film of eggs which we then interpreted as Rana clamitans. Besides a Rana
clamitans male was in this pool. This happened eight days before the dis-
covery of the small tadpoles. We now label the photo. Rana clamitans —
R. heckscheri (?), They may have been the latter. One would expect it to
lay a film of many eggs. In 1923 (p. 406) we placed the tadpole under those
frogs which laid their eggs in films. It was a mistake to place them so posi-
tively. They should have been placed in a category of "eggs unknown". An
estimate of the eggs of the two ovaries of the unspent female (123 mm. long)
is 8000-6000. This complement is greater than that of R. clamitans but no
more than that of Raiia grylio or Rana catesbeiana. The an mal pole is black;
the vegetative pole is white. The vitellus in diameter approximately ranges
from 1.5-2.0 mm. Not enough material is at hand to compare eggs carefully
with the above species. Mr. 0. C. Van Hyning reports their eggs and adults
from Georgia in 1930.

MATURE TADPOLE

In 1923 (p. 406) we characterized the tadpole thus: ''Rana sp. — black-
rimmed and black-banded tail, bluish venter, dorsum with greenish yellow
flecks."

Color description from life {July 21, 1922). Body dark greenish-olive or
ohve, finely covered with pale green-yellow or pale greenish-yellow flecks or
spots on the dorsum. On venter they are vinaceous fawn, vinaceous cinnamon
or orange vinaceous. Just back of angle of mouth in a mature tadpole and on
the venter is a clump of 4 to 6 much larger spots. Lower belly pale forget-me-
not blue to upper belly and breast jay-blue. Chapman's blue or grayish
violaceous blue. Spots of black become thicker on lower belly and at times
almost touch or make patches of color. Lateral line pores very prominent on
the head and body.

Ventral half of muscular part of the tail light salmon orange or apricot
buff or vinaceous cinnamon or ochraceous salmon. Upper half of the muscular
part with a black band on caudal two-thirds and more or less merged into
body color at its basal third. The black bandlike effect is produced by oblique

Rana heckscheri 391

bars of black where the myocommas are. These overshadow the intervening
body color. Whole rim of tail or edge of crests black, least just in front of
the vent.

Iris rim above and below orange cinnamon, tawny vinaceous, tawny or
orange rufous or better vinaceous rufous. Iris rim in front and behind
pupil black.

General appearance. Tadpole quite large (97 mm.), usually black of body,
and the most striking of all our (U. S.) Rana tadpoles. Belly pigmented so
intestine does not show through in preserved specimens. Tail elongate, tip
acuminate; dorsal crest not as wide as musculature width; not much different
from lower crest and not extending on body beyond the vertical through the
buds of the hind limbs. Spiracle sinistral, just visible from dorsal aspect,
divided backward and somewhat obhquely upwards. Spiracle usually with a
distinct semicircular impression on body and opening. Opening, leaving ex-
posed an elliptical or hemispherical patch on body. Spiracle clearly below
lateral axis (musculature axis). Eye on or just above lateral axis but in dorsal
aspect nearer lateral outline than mid-dorsal axis. Anus dextral opening at
edge of ventral crest.

Muciferous crypts distinct, white: a short dorsal row of a few pores on
either side of middle line of the back from the dorsal crest forward; from above
the middle Hne of insertion of tail musculature on body to a short distance
behind eye a prominent dorsolateral row; apparently resumed behind eye
after an interval and continued as supraorbital and infraorbital hues to above
and below the nostril; another lateral row from above insertion of hind legs to
gill region where a ventral commissure goes across to the row of the other side.
A third of the distance across the ventral branchial region a branch from the
commissure goes outwards and forwards along the jaw region almost to
the mouth.

Mouth parts. Teeth 2/3 or 3/3. Edge of upper labium greater than
length of upper horny beak and fringed by a continuous row of teeth. Some-
times this fringe is broken up as in figure. In either corner, beneath this fringe
is a short row of teeth about one-fourth to one-third of the length of the upper
fringe. The outer end of this second row never reaches outward beyond the
first fringe. Median space between these second rows of teeth, one to one and
one-half times the length of either lateral series of the second row. In some
median-sized tadpoles the space may be greater and the second row much
shorter or rarely absent. The third upper row very short, frequently absent
in young and medium-aged tadpoles. From above the end of the first upper
labial row of teeth to beneath the end of the third lower labial row are two or
three irregular rows of papillae which are continued across lower labium's
edge as one serrate row. The third labial row longer than this single row of
papillae equal to length of horny beak but 1/4-1/5 shorter than first and
second rows which extend beyond the ends of the horny beaks. The first row
is continuous or broken in the middle.

Measurements. Length of body (32.0-41.5 mm.) in tail (50-57.5 mm.)
1. 4-1. 85, average 1.625. Width of body (15.0^22.5 mm.) in its own length

392 Frogs of the Okefinokee Swamp

1.4-2.4, average 1.8. Depth (13-20 mm.) of body .9-1.6 in its own width,
average 1.14, rarely greater than body. Depth of body 1.8-2.46 in body
width, average 2.08. Depth of tail (14-18 mm.) in length of tail 2.6-4.6,
average 3.2. Depth of tail .8-1.3 in body depth, average 1.02. Muscular
part (9-1 1 mm.) of tail in its own tail depth 1.45-2.0, average 1.72. Spiracle
.86-1.2 nearer vent than snout, average i.o, i.e., about equidistant in general;
spiracle to snout (17-22 mm.) and spiracle to vent or base of hind legs (17-23
mm.). Spiracle to eye (8-12 mm.) in eye to snout (8-1 1 mm.) .85-1.2,
average .99, i.e. eye to snout and spiracle to eye usually equidistant. Nostril
to eye (4.0-6.0 mm.) equals the distance from nostril to snout (3.5-6.0) mm.
Mouth (3.0-7.0 mm.) usually i. 0-1.5 greater than the internasal space (3.0-
6.5 mm.), average 1.2. Mouth contained 1. 1-2.0 (average 1.46) in interorbital
distance (5.0-10 mm.). Internasal space contained in interorbital space 1.4-
2.0, average 1.75.

The dimensions of the largest tadpole of the 1922 collection are:

mm.

mm.

Total length

95-0

Spiracle to vent

23.0

Body length

41-5

Spiracle to eye

II-5

Body depth

17.0

Eye to snout

10. 0

Body width

17.0

Eye to nostril

6.0

Tail length

53-5

Nostril to snout

6.0

Tail depth

16.0

Mouth

6.5

Musculature

of tail

II .0

Interorbital distance

10. 0

Spiracle to snout

21.0

Internasal distance

5-0

In 1928 we took one 95 mm., one 96 mm. and one 97 mm. in total length.

General remarks. On June 16, 19 12, at the Fargo, Ga., heronry amongst a
swampy tangle of buttonbush Cephalanthus occidentalis, "hurrah bushes"
{Leucothoe raceynosa) and "lather leaf" {Clethra alnifolia) the author found
some black tadpoles with yellowish white crossbands and surmised that they
were the tadpoles of R. grylio. That was a mistake.

In 192 1 when I returned from the swamp three tadpoles were referred to
me from the U. S. Bureau of Fisheries. They were unlike anything we had
seen before. Our reply was solely as to the identity of the material. The
correspondent, a doctor from Savannah, Georgia, was concerned to know if
albinism was common in tadpoles. Some of them were albinos and some
normal. A week or so later, August 7, 192 1, Mr. Francis Harper and Marion
Lee found a small tadpole in the St. Mary's River, Baker County, Florida,
about 10 miles south of Moniac, Georgia. On August 16, 192 1, at Camp
Pinckney (3 miles southeast of Folkston, Ga.) St. Mary's River, they secured
three (Nos. 32-34) more larger tadpoles. These and the Savannah tadpoles
were at hand when we surmised they might be Gopher Frog {R. aesopus) or
Sphagnum Frog {R. virgatipes) tadpoles. They are of neither species.

In 1922 on July 17 at Thompson landing (south of Folkston) St. Mary's
River in a cut off overflow pool we found in the water almost a pure culture of
a small black tadpole with a gold and white transverse band like those of June
16, 1912.

Rana heckscheri 393

My journal reads thus: "In one cut off pool in a water course (which now
is a succession of separated pools) we hauled the seine. It was covered with a
wriggly mass which at first looked like water beetles to Miles (Dr. M. D.
Pirnie). I must confess I would have seen them the same way if I had not
seen them before. The tails are transparent and were hardly in evidence.
They have a band across the back. In another cut-off from the river proper
(St. Mary's River) in shallow water they were in immense numbers and pre-
sented a very beautiful sight in sunlit situations with their dark bodies and
transverse bands. Were it not for the transverse bands, they would look like
toad tadpoles. Then I provisionally placed them with the green tree frog,
Hyla cinerea, which sometimes has somewhat the same appearance."

Three days later, July 20, we started for Jacksonville, Florida. "At
Callahan, Florida, (just north) near a large concrete bridge for the Dixie
Highway, a car was stuck on the smaller bridge to the west in the detour and
we had to wait. In the areas beside the new Dixie Highway were shallow
ponds or overflowed areas. These were tributary to Alligator Swamp which
in turn is a part of Mills Swamp (U. S. Geol Survey Sheet, Hilliard). At first
I saw a few cross-banded forms of the Thompson's landing sort which I took
to be Hyla cinerea. Now I suspect they are probably R. aesopus or R.
virgatipes, probably the former. We collected a few and went on. They are
very conspicuous with black rimmed crests and black bands on upper half of
tail musculature and a light color on lower musculature. In one-third grown
ones the cross bands show through faintly. When full grown it disappears."

On July 21, on our return from Jacksonville "we stopped at Callahan
(Alligator Swamp). The tadpoles of July 20 were abundant. They travel
in big schools as no other big tadpoles do. They remind me of a school of
mature Bufo tadpoles. Once in a while amongst the fair sized ones were
monsters almost as big as a bullfrog tadpole. And these monsters have no
suggestion of hind leg buds. Does this species winter over one or two years
as a tadpole?"

We checked up our Thompson Landing (Georgia) and Alligator Swamp
(Florida) material and found them all of the same species. We later found
they could not be tadpoles of Rana aesopus or Rana virgatipes.

On June 8, 1928, we went "to the type locality for Rana heckscheri. Saw
plenty of tadpoles amongst vegetation under the Dixie Highway bridge.
Captured about 12 transformed individuals. Can approach them when head
is out of water and can catch them by hand. Or they may rest on mudflats
where we also secured some. Many are with 4 legs and a long tail. . . . The
transformed frogs are somewhat like greenfrogs but with more intensified
spottings on the belly. These as transformed frogs and as adults have no
lateral folds."

One June 9 "at Mr. Davis's brick yard in the large clay pool in the shade
of the trees amongst Polygonum hydropiper were plenty of these tadpoles and
transformed frogs near or at the edge of the water. Here in the shade they
hopped about in amongst the weeds. They hid amongst vegetation, under
boards and sticks or finally hopped into the water. There were a few bigger

394 Frogs of the Okefinokee Swamp

ones with them but they were more wary. Countless tadpoles are in this
immense clay hole. Sometimes the tadpoles are so thick that one can reach
in with his hands and catch them."

"On the east side saw prodigious schools of little black tadpoles with
yellowish cross bands on the body. Amongst them are some bigger ones,
about one-half inch in body length. This means two lots or breeding periods
this year."

Oftentimes these schools would mass like a school of young catfish. At
times one tadpole would lead out and the whole school would go in single file
or the head and tail of the line might meet and all merge into mass formation
again. Frequently smaller schools of the bigger size would be segregated
from the smaller ones. All kinds of formations were made and the tadpoles
are striking, either young or old. These pools have waterlilies, Sagittaria,
water hyacinth, Hydrocotyle, etc.

LARVAL PERIOD

If I did not know that in the South there are some Rana catesbeiana
which transform in one year or in the North Rana septentrionalis which some-
times spend two winters in the tadpole stage, I would feel inclined to believe
this species always spent two winters in the tadpole stage. The largest tad-
poles reach about loo mm. When we caught them in July (2 ist) or in August
(i8th), there were at least two or three distinctive sizes of tadpoles. The
largest ones were scarce. Doubtless the vast bulk of tadpoles at this time
July 2i-August 18 were one year olds or newly hatched tadpoles.

Our tentative conclusions above of 1922 in general express the situation
as we see it in 1928. In 1928 we took tadpoles which drop into two groups
namely, small and very large. The first of the clay pit of Callahan, Florida,
are manifestly of the first year and appear to fall into three sizes: 10-15 mm.
(body 7 mm.), 20-25 mm. (body 10 mm.), 30-35 mm. (body 12-15 mm.).
Apparently this means three different breedings in the spring. The other
group are the large transforming or near transforming tadpoles. Certainly
it seems as if two years might be needed to arrive at the maximum size yet
we found in 1922 and 1928 no intermediate tadpoles between the very large
ones (75-100 mm.) and the 30-35 mm. sizes which might imply a shorter
larval period.

In 192 1 the three secured from the U. S. Bureau of Fisheries from Savan-
nah, Ga., measured 44, 60 mm. albinos and 70 mm. normal. The three taken
by Mr. Harper and Marion Lee measured 83, 88 and 90 mm. respectively.
Except for one of these last the hind legs were pigmy in size. This material
does not seriously mihtate against two years for larval life though one year
may be all.

TRANSFORMATION

Period. On July 20, 21, 1922, and August 18, 1922, there were few of the
large tadpoles. Doubtless transformation comes mainly before July. The
few mature tadpoles of these dates were quite clearly stragglers. They might
not have transformed before September loth or 15th.

Rana heckscheri 395

The paragraph before was more in the nature of negative evidence and
conjecture but on June 8-10, 1928, countless mature tadpoles were approach-
ing transformation. Our series of 89 frogs transformed and transforming
could have been materially increased had we wished. Probably as in Rana
catesbeiana, R. damitans and some of the other large bodied tadpoles trans-
formation is largely accomplished by the end of June. A few of the trans-
formed ones (40-49 mm.) might indicate transformations in May, and some
of the mature, about-to-transform tadpoles might not transform before July
I or possibly July 15. In other words before the period in which we observed
them in 1922.

Size. In 1922 we held that "some of the largest tadpoles with large bodies
and well developed hind legs had body lengths of 32-41.5 mm. and the trans-
formation size probably falls within this range." On June 8 and 9, 1928, we
took 89 transformed or transforming frogs: — transformed (41) or nearly so
with short tails (3-29 mm.) (48 specimens). Almost invariably if there was
an appreciable stub the mouth was not the full-developed frog mouth. We
can properly say that the range of size reaches from 31-49 mm. The 89
start with three at 30 mm., reaches the mode at 33 mm. with 22 specimens,
and almost ceases with three at 38 mm. The record onwards is i at 39 mm.,
I at 41 mm., one at 43 mm., 44 mm. and 45 mm., 3 at 46 mm., i at 48 mm,
and I at 49 mm. We thus see there are only nine of the 89 which are 40
mm. or more. The normal range is from 30-39 mm. yet there is clearly
another smaller group with a mode about 46 mm. The average of the 30-39
mm. is 34 mm. or of the whole 89 specimens from 30-49 mm. is 35.5 mm,

GROWTH

On August 18 we captured at least four different sizes of frogs some (8-10
frogs). The same evening we later lost all but three of them. These are 56,
82 and 95 mm. respectively. The range of body length of seven large tadpoles
with hind legs well started is 36-40, average 38, mode 39 mm. It is true we
have given 32-40 or 41 for transformation size but the bulk of it probably
comes within the narrower range of 36-40. This would give us 32-41 for
transformation. The 56 mm. specimen may be two years old, the 82 mm.
specimen a four year old and the 95 mm. specimen a five year old. They were
all taken at the same time and there was another group of sizes between
56-82 mm. amongst the 8-10 specimens originally captured. (1922).

The conclusions above of 1922 are hardly correct for the 56 mm. specimen
— the smallest frog we then had. If transformation sizes of June 8 and 9,
1928, vary from 30-49 mm. the 56 mm. can not be two year olds. It seems
quite likely that some of the frogs which are almost transformed may retain
their tails some time thereafter or that transformation may at times be
delayed beyond the normal 30-39 mm. The premise of 1922 that 32-41 mm.
was the probable transformation size is not far from the normal 30-39 of
1928. An unusual 49 mm. transformed frog would have to grow only 7 mm,
to attain 56 mm. which can hardly be a two year old.

396 Frogs of the Okefinokee Sivamp

In 1928 we collected many intermediate sizes between transformation
and 82 mm., but they were mislaid. These with the last 5 live intermediates
of 1922, might have outlived the possible growth groups. With our meager
material we might compare it with the possibly comparable Raria grylio.
We might postulate: 30-49 mm., average 35.5 mm., transformation; 56 mm.,
I year old; 82 mm., 2 year old; 91, 95, 95, 97 mm. cf s, 102 mm. 9 , 3 year
olds; 113 mm. cf, 4 year old; 125 mm. 9 , 5 year old.

ENEMIES

On June 8, 1928 we found a large southern water snake (Natrix sipedon
fasciata) on the mud flats where transformed frogs were or at times it was
coursing in water where countless tadpoles were. The pied water snake is
also an enemy of these tadpoles and young frogs. On June 9, 1928, in the
clay hole where the youngest tadpoles were numerous large mouthed black
bass lived. These doubtless feed on the tadpoles. At the same time Florida
grackles were apparently feeding on the transformed frogs along the edges of
the pond.

At the type locahty we took one transformed frog with all of left leg gone
below femur. Another had the left leg completely severed and the toes clean
cut from the right foot. In this hole we took Sternotherus minor. The above
work is quite clearly the work of turtles or possibly fish.

AFFINITIES

The original diagnosis is pertinent in this instance.

Diagnosis. Like Rana grylio and Rana catesbeiana, it has no dorso-lateral
fold and no phalanx of the fourth toe is totally free of web; third toe in 56 mm.
specimen is i to 3 mm. shorter in Rana heckscheri than in the other two species
of bullfrog or 3 to 6 mm. shorter in 82 mm. specimen or 6 to 9 mm. shorter in
95 mm. specimen; third toe 3.8 (95 mm.)-3.56 (82 mm.)-3.3 (56 mm.) in
length (snout to vent) in R. heckscheri while 2.7 to 3.1 (95-56 mm.) in R.
grylio and R. catesbeiana; third toe 1.6-1.7 in fourth toe in R. heckscheri
while 1. 2 -1. 5 in fourth toe in the other two species; first finger decidedly
longer than second, while in the other two species it is usually shorter or
sometimes equal; first, second, third and fifth toes shorter than corresponding
toes of R. grylio and R. catesbeiana; fourth finger 8.6 (95 mm.)-8.2 (82 mm.)-
8.0 (56 mm.) in length (snout to vent) while 6.0 to 6.3 (95 mm.)-6.8 to 7.4
(82 mm.)-5.6 to 7.1 (56 mm.) in the other two specimens; internasal space
less than upper eyelid width, i. 07-1. 43 in it while .85-1.0 in R. grylio and R.
catesbeiana; tympanum in males is proportionately greatest in R. grylio,
somewhat smaller in R. catesbeiana and R. clamitans and smallest in R.
heckscheri; intertympanic width of 95 mm. male in length (snout to vent)
4.52 in R. heckscheri, 5.43 in R. clamitans and 6.3 in R. grylio (R. catesbeiana
males of 95 mm. have tympanum poorly developed, but a 136 mm. has it 4.85) ;
in general, intertympanic width broadest in R. heckscheri and R. catesbeiana
and narrowest in R. grylio; distance from the rear corner of the eye to the

Rana septentrionalis 397

same corner of the other eye much greater than the intertympanic width in
R. grylio, somewhat greater in R. clamitans, about equal in R. catesheiana
and equal in R. heckscheri, i.e., in the males.

In the field at night we mistook the first one for a green frog until another
specimen called, but in our original description we called it a new bullfrog.
More material and more life history clues are needed to establish its place
in the Rana catesheiana- Rana clamitans series.

BIBLIOGRAPHY

1923 Wright, A.H. The Anat. Rerord, Vol. 24, No. 6, p. 406 (as Rana sp.).
X924 . Proc. Biol. Soc. Washington, Vol. 37, pp. 141-152.

Rana septentrionalis Baird

(PI. Ill, Fig. i; IX, Fig. i; XI, Fig. 11; XIV, Fig. 3; XVI, Figs. 9, 10; XXXIX)

COMMON NAMES
Mink Frog. Northern Frog. Hoosier Frog. Rocky ^Mountain Frog.

RANGE

Check list. Type locahty: Northern Minnesota. Range: Northern New
England and Northern New York, west through Michigan to Minnesota,
Canada, northward to Hudson Bay. Stejneger & Barbour Check List

1923, P- 37.

Supplementary records. Miss Dickerson relying on Cope says "It is re-
ported from the Adirondack Mountains, from Lucknow, Ontario, and Fort
Ripley, Minnesota, and from Moose River and the Hudson Bay region.
Strictly speaking, Madrid and Garrison's Creek, N. Y., are not Adirondacks
proper but Paulmier found one on Black Lake in the Fulton Chain. On June
13 and 14 A. H. and A. A. Wright found the species from White Lake to Old
Forge, N. Y. On June 28, Messrs. S. C. Bishop, C. K. Sibley, C. R. Crosby,
M. D. Leonard, P. W. Claassen and L. West found it at Hart or Clear Lake
(Mt. Marcy region). On July 7, Mr. C. W. Leister also recorded it there as
did Mr. and Mrs G. B. Upton from July 7-12. On July 13-15 Dr. S. C.
Bishop studied it in this place. Mr. C. W. Leister has also seen it in Connery
Pond, about four miles northeast of Lake Placid. G. A. Boulenger (1920, p.
424) in his range Southern Canada and New York to Montana and Utah
omits Pope's records 1914-1918 from Maine, yet No. 8 in his measurements
from Eustis, Maine, must be a Pope specimen. With Pope's discoveries of
this species in Frankhn County, Me., with the general distribution of this
species in the Adirondacks, search at the right time ought to reveal the species
in New Hampshire, Vermont and possibly in the Catskill Mts., N. Y. and
doubtless in some of the swamp area around Oswego, N. Y., or north of
Auburn, N. Y. Nash, (p. 10) holds that it extends in Ontario from "Bruce
County and northward and westward, not common." He doubtless is

398 Frogs of the Okefinokee Swamp

considering the Lucknow record. The parallel of Lucknow 44° about repre-
sents the more common southern limit of its range in Ontario and New York
but records are already at hand as low as 43°, 30' in New York State.

In Ontario, it is knows from Haliburton to Bruce Counties and doubtless
it is in the country east of HaHburton Co., Ontario to Madrid, N. Y.

Two years after the above paragraphs were written, we examined the U. S.
National Museum series. We find a Vermont record: Newport, Clyde River,
Evermann & Bean, July 23, 1894 (U. S. N, M. 39813-4). A New Hampshire
accession of 4 specimens (U. S. N. M. Nos. 36463-6) from First Connecticut
Lake, N. H., July 21, 1904, is made by the Bureau of Fisheries. This gives
a continuous stretch of states or provinces from Maine to Manitoba. To me
the most interesting recent record is the three series of accessions (U. S. N. M.
Nos. 28450; 28336-46; 28354-58) of Mrs. E. P. Miller and Mr. Gerrit S.
Miller Jr., from Peterboro, Madison County, N. Y. These are the farthest
south in New York State unless it be E. A. Mearn's record for the Catskills
(Am. Mus. Bull. Vol. 10, 1898, p. 325). These Peterboro records strengthen
the probability of Rana septentrionalis being in the Catskills.

GENERAL APPEARANCE

This frog is a small representative of the bullfrog-green frog group. The
sides are heavily mottled, the rear of the femur heavily reticulated, the back
may be almost uniform, may be mottled with large dark areas set off by a
tracery of light lines around or amongst them, or the back may be spotted
with widely separated spots. Sometimes the forward part of the back is
uniform and the rear part spotted. The legs are spotted or with a few bars.
The mottling on the femur is suggestive of the R. virgatipes-R. grylio group.
In young specimens the sides may be speckled and throats mottled.

MEASUREMENTS

(Recent Material)

Head to angle of mouth 1.2 (36 mm.) — 1.07 (44 mm.) — 1.24 (56 mm.cf) —
I.I (56 mm. 9) — 1.25 (66mm.cf ) — 1.13 (66 mm. 9 ) in width of head; head to
rear of tympanum .92-.94-.9-.95-1.04-.93 in width of head; head to angle of
mouth 3.6-3.13-3.4-3.3-3.3-3.47 in length of body; head to rear of tympanum
2.66-2.75-2.54-2.9-2.75-2.87 in length of body; snout 1.16-1.14-1.11-1.18-
1.1-1.15 in first finger; snout 1.16-1.14-1.4-1.31-1.4-1-16 in fourth finger;
snout 1.25-1.28-1.0-1.125-1.4-1.05 in first toe; eye i. 5-1. 27-1. 13-1. 45-1. i in
snout; eye .75-.81-1.13-1.0-1.06-.92 in tympanum; eye i. 75-1. 45-1. 26-1. 74-
1.3-1.7 in first finger; tympanum 3. 3-2. 44-1. 3-2. oi-i. 4-2. 33 in intertympanic
width; tympanum 2.0-1.6-1.0-1.45-1.11-1.6 in snout; internasal width i.o-
.75-.8-.9-.91-1.0 in upper eyelid width; interorbital width 1.4-.86-1.8-2.25-
1. 83-1. 66 in upper eyelid width; interorbital width i. 4-1. 14-2. 2-2. 5-2. 0-1.66
in internasal width; interorbital width 4.0-3.14-4.4-6.0-4.0-4.56 in intertym-
panic width.

Forelimb: Forelimb i. 9-1. 9-1. 8-2. 1-2. 3-2. 3 in length of body; forehmb
3.1-2.7-2.66-2.9-3.3-3.2 in hind limb; first finger i. 3-1. 4-1. 47-1. 3-1. 66-1. 3

Plate XXXIX

Mink frog (Rana septentriojialis)

1 . Male, collected by S. C. Bishop at Hart (Clear) Lake, Adirondack Mts., N. Y.

July 5, 1923. Lateral aspect, x i.o.

2. Female, collected by S. C. Bishop at Hart (Clear) Lake, Adirondack Mts., N. Y.

July 5, 1923. Ventral aspect, x i.o.

3. Beaver dam, Hellgate pond, Onekio, N. Y. June 12, 1923.

4. Egg mass, photographed by S. C. Bishop, Hart (Clear) Lake, Adirondack Mts.,

N. Y. X 0.5.

5. Transforming frog with short tail, Onekio, X. Y. July 4, 1923. Dorsal aspect.

X I.o.

6. Adult, photographed by C. \V. Leister, Hart Lake, Essex Co., N. Y. July 7,

1923. X 0.6.

Rana septentrionalis 399

in third finger; second finger i. 3-1. 4-1. 33-1. 4-1. 6-1. 3 in third finger; second
finger 1.0-1.0-1.0-.95-1.0-.91 in first finger; third finger 1.66-1.3-.93-1.2-
1.05-1,0 in second toe; fourth finger 1.3-1.45-1.16-1.2-1.3-1.3 in third finger;
fourth finger 1.07-1.15-.70-.85-.8-.9 in first toe; internasal width 2.0-2.0-
1. 74-1. 9-1. 75-2. 2 in first finger; internasal width 2. 0-2.0-1. 9-1. 8-1. 83-2. 2 in
second finger; internasal width 2.6-2.87-2.5-2.5-2.9-2.9 in third finger; inter-
nasal width 2.0-2.0-2.05-2,1-2.25-2.2 in fourth finger.

Hindlimh: length i. 64-1. 46-1. 45-1. 3 7-1. 45-1. 3 in hind limb; tibia 2.0-
2.0-2.0-2.2-2,2-2,2 inlength; tibia3. 2-2. 93-3. 0-3. 0-3. 2-3. 03 in hindlimb;tibia
1. 05-1. 07-1. 08-1. 03-1. 0-.93 in forelimb; tibia 1.08-1,13-1,08-1.1-1,03-1.03
in hind foot (without tarsus); first toe 1.3-1.55-1.53-1.55-1.63-1,45 in second
toe; first toe 2,0-1,9-2,6-2,1-2,04-2.3 in third toe; first toe 2.66-2.66-3.5-
3. 1-2.7-3. 1 ill fourth toe; first toe 1.9-2.0-2.3-2. 16-2.2-2. i in fifth toe;
second toe 1.5-1.2-1.7-1,4-1,25-1.6 in third toe; second toe 2.0-1,8-2.3-2,0-
1.7-2,1 in fourth toe; second toe 1,4-1,3-1,5-1,3-1,4-1,5 in fifth toe; third
toe 1,3-1,47-1.36-1.46-1.4-1.35 in fourth toe; third toe o,93-i,o5-,9-i,o3
i.i3-,93 in fifth toe; fourth toe 1,3-1,3-1,3-1.4-1.53-1.34 in hind foot (with
tarsus); fourth toe ,925-,9-,9i-,9i-.96-.96 in tibia; fourth toe ,95-94-
i.oi-.94-,96-,9 in forelimb; fifth toe 1,38-1.4-1,5-1,43-1,2-1.4 in fourth toe;
internasal width 2. 14-2. 25-1. 54-1. 80-1. 83-2.0 in first toe; internasal width
3.0-3.5-2.3-2.8-3.0-2.9 in second toe; internasal width 4.4-4.25-4.0-3.8-
3.75-4.6 in third toe; internasal width 5.7-6.25-5.4-5.6-5.1-6.2 in fourth toe;
internasal width 4.1-4.5-3.4-3.9-4.25-4.3 in fifth toe.

HABITAT

Garnier (p. 945) pronounces this species a river frog. "It inhabits spring
creeks and rivers, but in lakes and ponds of the purest water I have never seen
it nor captured a single specimen." "It is . , , emphatically a river frog.
It is never seen in fields nor woods lurking. ..." Pope (191 5, Copeia No 16)
records "one from a wood road, which passed through a sphagnum bog,"
''Rana septentrionalis common along the banks of Ciss Stream, which flows
from Round Pond into Caucmogomoc Lake, Piscataquis Co. (Me.) . , , , They
were found on the marshy banks of the stream where bull frogs {Rana cates-
heiana) were abundant," Mr, Arthur H. Norton (Pope, Copeia 1915, No,
16) observed it "in VanBuren and Caswell, (Me.), August 15-16, 1914. It
was found to be numerous in two dead water ponds in Caswell, locally called
Mud and Guard Ponds. They were more conspicuous than Rana clamata,
inhabiting similar places, i.e., tussocks of sedges on the edges and the beds
of lily pads near the shore. At Van Buren they were found in ditches by the
railroad track and were common," Pope concludes the mink frog is found in
the same localities as R. catesbeiana and R. clamitans, not tending to replace
either species. Later Pope (Copeia Dec. 31, 19 18, p. 96) "found it abundant
at Tim Pond, Eustis, Franklin Co., Me. It occurred all around the pond;
along rocky, wave-beaten shores, in marshes, and in the outlet close to the
lake, but never far from water."

400 Frogs of the Okefinokee Swatnp

In 19 13 (Copeia October 15, 19 15, pp. 46-48). "On July 7 in peaty lake
with a clear sphagnum border we found several R. septentrionalis.'" On July
1 6th and 17th we examined the place closely. "All along the north edge of
the lake were white water lilies, yellow spatterdocks and water shields.
These three made a perfect carpet on the water's surface. On these plants
during the day the frogs rested." "In the outlet to Otter Lake (Ten Mile
Creek between Lake of Bays and Otter Lake) we found them common,
July 24, on muddy bottoms where water lilies were abundant. In the same
kind of situation they occurred on Porridge Lake, July 28. Another habitat
we discovered August 31 was a beaver lake where Cassandra and all the
associated heath-like plants grew. Finally, in Fletcher Lake, September i
we found them in the shallow, sandy shores amongst pipeworts (Eriocaulon
articulatum).'' "This species cannot be called solely a river species; . . ."

To the last conclusion I still adhere. In general we have found it most
commonly in lakes or ponds or if streams these are usually inlets or outlets
of ponds or lakes. On June 13-14, 1923, we found it more or less along the
whole Fulton chain of lakes, Adirondack Mts. from White Lake to Seventh
Lake. Sometimes a few might be in temporary pools beside the state road
where they must have travelled over land to reach these places. All in all,
it is distinctly an aquatic species. On June 14, 1923, we found most of our
frogs at Hellgate Ponds, Onekio, N Y. Here in driftwood pools below a
beaver dam we took several R. damitans, R. catesbeiana and R. septentrionalis.
Later took several more mink frogs in muddy pools among granite boulders.

On July 9, and July 26, 1900, Mrs. E. P. Miller and Gerrit S. Miller Jr.,
recorded these frogs in Peterboro Lake and Peterboro Mill pond on floating
Nymphaea leaves, the same habitat in which Bishop in the Adirondacks and
we in Algonquin Park found them.

FIRST APPEARANCE

None of our records are in the spring and the earliest record of the year is
Garnier's statement "It makes its appearance in April" at Lucknow, Ontario.

GENERAL HABITS

Variations in color. This topic is not fully treated or considered. Some
scattering notes on some U. S. National Museum specimens are:

No. 39813. Male 58 mm. Dark spotted on upper head. Green from

tympanum to snout in sharp contrast to top of head. Green involves

edge of eye except upper eyelid.
No. 39814. Just transformed 31.5 mm. Sometimes dark spots have

light centers.
No. 39792. 55 mm. 9 . Almost no spots on upper parts but mottling

on sides, rear of femur and legs barred.
Nos. 28354-58. 51 mm. cf, 66 mm. ripe 9, 57 mm. cT, 56 mm. cf,

66 mm. 9 . These have large spots on back, some light encircled,

some not.

Rana seyientrionalis 401

Nos. 36096-99. 62 mm. 6^, 62 mm. 9 , 57 mm. cf , 56 mm. cf . These
are all with distinct and separate spots on back.

No. 3632. 41 mm. Speckly on sides.

Nos. 28450-56. 41 mm. -48 mm. 9, backs almost uniform.

No. 3437. 53 mm. c?'. Spots few on rear of back only. Legs much
spotted.
72 mm. 9 . Forward part of body less spotted.

No. 5245. 58 mm. 9 , 67 mm. 9 . In general mottled with large
dark areas and a tracery of light around them.
This species is quite common in more northern parts. Except at its short
croaking period it is generally passed by as a small bullfrog or green frog,
and as a consequence is pronounced "not common," "rare." It is called
northern frog from its specific name and from its northern distribution, — a
not inappropriate name except it is not individualistic. Gamier hke others
subsequently noted its secretions. He writes (1883, p. 945) "If taken in the
hand it emits a strong odor of musk and garlic, or more properly the disagree-
able scent of the mink; this is sufficiently powerful to adhere to the hand for
a time, but soon passes away." (p. 948) "The tadpole has the odor of the
frog, though not so strong; ..." In 19 13 I thought the term "mink frog
very apropos for this species, at least for Ontario specimens. ..." I have
since noticed some have it more than others and possibly the males have it
strongest at the breeding season. At times some Rana septentrionaUs have
the scent very faint and do not always respond with the secretion flow when
rubbed. All in all, this is the best name suggested. One day I put several
in a glass jar with several meadow frogs, some R. virgatipes and two green
frogs. Almost all were dead on the following morning including the two mink
frogs. Did the secretions cause it as some times happens with R. palustris*
The terms "hoosier frog," "Rocky Mountain frog" seem inappropriate.

Of its general habits Gamier writes (pp. 940, 948) "In summer they may
often be seen with the head and a bit of the back out of water, resting among
plants on the borders of streams, and where the Potamogeton is in bunches,
or the Ranunculus is in beds, the herpetologist may likely secure his speci-
mens." "But if the frog once disappears, he generally keeps from view till
all probable danger is past. It is useless to expect it to reappear at the spot
it left, as it dives several yards, it may be, rods, before it stops. I have
occasionally waited half an hour or more, watching one that has so dived
beside a stone, in the current or otherwise. Perhaps they may have been so
really frightened and the feeling of fear may have remained, or they may
have followed some law of nature implanted with them in keeping concealed
for such a protracted period. Occasionally I have heard their notes after
they were secured and in my collecting case; but then it seemed truly a note
of distress, and was in a different tone and key from that rarely heard in the
open stream."

The experience of several subsequent collectors prove them not hard to
capture at all at the breeding period and Mr. Simpson's capture of 30-40 at
one time in August from lily pads with an insect net hardly accords with
Garnier's results. Possibly stream or river specimens may be harder to secure.

402 Frogs of the Okefinokee Swamp

VOICE

Garnier (1883, pp. 949, 950) writes ''that (croaking) of the mink frogs
is a rapid squeaking croak almost Hke the notes uttered by a toad when seized,
with the finger and thumb, by its arm pits. I have since heard the same love
cry late in the evening, on the banks of the stream and have well recognized
its peculiarly sharp ringing croak." "Occasionally I have heard their notes
after they were secured and in my collecting case; but then it seemed truly
a note of distress, and was in a different tone and key from that rarely heard
on the open stream. . . . There are no 'chant amour' or love notes, in spring"
(pp. 947, 948). In 1913 near Dorset, Ontario, "from July 7-14, we heard at
night along the shore of Otter Lake the pecuhar note which later proved
the croak of the mink frog. On the 15th of July at 10:00 p.m. we heard
several frogs and started with flashlight for Peat Lake where the species was
in chorus. The air temperature ranged from 52° to 55° F., but the water of
Otter Lake at its surface registered 69° F. . . . After July 15th and i6th we
heard no more choruses. . . . (This species) has a 'chant amour' which at
chorus season can be heard one-third to one-half of a mile away; ..." (Wright
1915, Copeia No 23, pp. 46, 47). In Maine, P. H. Pope writes of it as follows:
"Through the kindness of Mr. Arthur H. Norton I am permitted to publish
the following data collected by him:

'At Mud Pond, Caswell, they were heard calling, and the same is true of
R. clamata.

'The call of R. septentrionalis was found to be similar in nature to that of
R. clamata, but differed strikingly in tone, being higher and slightly metallic,
resembhng closely the sound produced by striking a long nail on the head
with a hammer in driving it into heavy timber.' " Pope (191 5, No. 16, p. 2).
Later Pope in a subsequent note (Copeia, 1918, p. 96) remarks "I also heard
the call of these frogs and find that Mr. A. H. Norton's description, quoted
in the paper mentioned above, fits it admirably. While keeping some of these
frogs in camp in a tin can I heard them give a slightly different call suggesting
the bubbling note of Rana pipiens but shriller and not quite so loud." F.
Harper (1926, p. 11) writes as follows of these frogs at Clear Lake, Adiron-
dacks: "The notes were heard occasionally by day during cloudy weather,
but perhaps more frequently by night. There are guttural, vigorous, and
rather rapid: Cuck-cuck-cuck-cuck.^^

From the previous published data, croaking has been recorded from
July 7-August 16. In 1923 some captives we took June 14 later croaked,
but we did not hear them in the field. On June 28 at Clear or Hart Lake,
Adirondack Lodge, N. Y., Messrs. S. C. Bishop, P. A. Claassen, C. R. Crosby,
M. D. Leonard, C. K. Sibley and L. West heard them. On July 7, 1923,
Mr. C. W. Leister heard them in the same place and also near Saranac
Lake. About the same time Mr. and Mrs. G. B. Upton also heard them at
Hart Lake. On July 14 Mr. S. C. Bishop found them in chorus at Hart
Lake. At Onekio, N. Y., Hellgate Pond on July 14 I heard many individual
croaks in the daytime.

Rana septcntn'onalis 403

On July 2 at Adirondack Lodge, Hart or Clear Lake (2155 ft.) Messrs.
S. C. Bishop and C. K. Sibley found mink frogs croaking at 8:30 a.m. in
broad sunlight, at mid-day and in the evening. "On July 13 in the same place
S. C. Bishop heard males croaking just at daybreak in the middle of the morn-
ing and rarely in the afternoon." On July 14 he records "Frogs croaked all
night. The male croaking expands the throat and sides of head below ears.
The common note is a cut-cut — with sometimes a 'burred' gh-r-r-r." On July
15 "dark and rainy. Only a few frogs croaking during the night and very
infrequently in the daylight. P.M. No croaking this period."

This species seem to have about the same sac development as R. palustris.
In some from Onekio, N. Y. it seemed that the head if viewed from dorsum
appears at croaking to have no perceptible lateral sacs. Viewed from below
the central throat swells out with a sac on either side of this central part.
This sac however appears to be just below corner of the mouth or directly
below the vertical of the tympanum but not directly below the ear or between
ear and shoulder .

From these we concluded it did not swell out as in a Cope's frog male.
A male Mr. Bishop brought me from Clear Lake has the lateral sacs extending
between tympanum and shoulder. The caudal margin of the lateral sac
beyond the rear vertical of the tympanum or even to the vertical of the rear
arm insertion. In this frog it is very apparent from the dorsum but not so
striking as in Cope's frog. This frog, as with other species of frog, can be
made to croak by seizing it just in front of its hind legs or better by laying
it on its back and by rubbing its belly. Then it will croak continuously.
Each individual croak in nature occupies about one second or less. If not
in chorus their croaks are not heard long distances when their associates,
green frogs or bull-frogs are calling. At Onekio, N. Y. we heard them. On July
14 we could hear the males croaking 60-100 yards away. In croaking one
or two frogs seemed to move forward in the water with every croak and the
body return to position immediately after each croak. The vocal sacs were
not very well revealed in the field. A frog usually gave two croaks, occa-
sionally one, occasionally 5 or 6. Its croak is different from that of the Cope's
or Carpenter frog though Norton makes a carpenter comparison in describing
the croak.

MATING

Male (From life June 18, 1923). On back buffy olive, light brownish
olive or olive; this same color becomes Isabella color on upper side of hind
legs, amongst this buffy olive, etc. are wavy lines or spots of oil yellow, yellow-
ish citrine; on the middle of the back are two irregular rows of black spots
often round, widely separated, at times elongate. Region of dorsum forward
of rear of tympana is without black spots but buffy olive with plenty of cosse
green or lettuce green interspersed. This color solid on the upper jaw to
level of nostril to tympanum. Above and below tympanum a line of courge
green. Just back of tympanum courge green or yellow green spots and lines
with black. On sides mixture of yellow-green, black and hazel (or tawny
olive or fawn). Tympanum tawny olive, buffy brown or Isabella color. Upper

404 Frogs of the Okefinokee Swamp

parts of fore legs citrine drab or buffy brown with black spots of fore edge
and rear edge distinct and small. Hind legs with vermiculation of buffy
olive, etc., and black on dorsum. On posterior thighs fine black and old
gold or olive lake. Rear thigh parts if viewed from venter suggest R. virga-
tipes somewhat and R. grylio. Throat of male sulphur yellow or pale green
yellow, rest of under parts seafoam yellow. Pupil rim green yellow; iris
finely spotted black and vinaceous tawny or xanthine orange. Thumb
somewhat enlarged.

(From life July I4, 1923). Upper jaw solid courge green, cosse green or
lettuce green; also this color more or less predominant on head back to
tympanum. This color less prominent in back because of large black spots
which are outlined by these greens. Amongst these black spots are smaller
snuff brown spots. On hind legs are sulphur yellow or in places aniline yellow
vermiculating the black of the thighs and bounding the black cross bars of
the lower legs. More or less longitudinal bar on thigh bister or brownish
olive. On sides mixture of black, green yellow or even yellow-green and
hazel (very slightly). Upper parts of fore legs yellowish oil green with black.
Throat maize yellow or sometimes baryta yellow while the rest of underparts
are light chalcedony yellow. Some of the largest males may have maize
yellow on entire under parts. Iris as in description of male of June 18.

Female {From life July I4, 1923). Several seem more spotted on under
part of hind legs than in males. Sometimes spots on either side of the throat.
Hazel of back more prominent and more hazel on sides and usually more
spotted on the head. Underneath they are white or whitish except for the
throat which may be pale chalcedony yellow or seafoam yellow.

Structural differences. Some of the differences are :

1 . Males have tympana larger than rarely equal to the eye while those
of the females are clearly smaller than or equal to the eye. For
example, extreme tympana may be as follows: cf 56 mm., tym-
pana 8 1/2 mm., eye 5.5 mm., or cf 56 mm., tympana 7.5 mm.,
eye 6.5 mm., or cf 54 mm., tympana 8.5 mm., eye 7.5 mm.
Occasionally the tympanum may reach 10 mm.

2. A pair taken at Hart Lake, Adirondack Lodge, July 14, 1923,
show the following differences in ventral coloration. The male's
throat has some maize yellow or baryta yellow while the rest of
the under parts are light chalcedony yellow; the female has a sea-
form yellow throat with some chartreuse yellow while the rest of
the under parts are white or whitish.

3. Vocal sacs (see Voice). In a 57 mm. cf from Peterboro, N. Y., a
little inclination toward a sac back of tympanum.

4. General coloration (see Mating coloration).

5 . Tympanic borders prominent in male. Old males are often with
the tympanic border standing out prominently from the surround-
ing body.

6. The males have the characteristic swollen thumb of our North
American Ranas.

Rana septentrionalis 405

7. Males probably do not breed until two years from transformation.
In our series we have no positive male (externally) before 50 mm.
In the U. S. National Museum series is one 48.5 mm. Of one speci-
men in our series, 46.5 mm., we record that it "begins slightly to
show maleness in tympana and thumbs."

8. In like fashion the females do not breed until two years from
transformation. In our series the first positive female is at 50 mm.
In the U.S. National Museum series we recorded in Miller's speci-
mens from Peterboro, N. Y. a 9 47 and a 9 48 mm. In our own
material we have one questionable female at 48 mm.

9. The range of size for positive breeding males is from 48.5-66 mm.,
for females from 50-72 mm.

Duration, night or day. The mating habits of this species still rest on few
observations. The period of mating begins in the latter part of June and
extends well into August. Garnier (1883) found eggs June 24 and July 30.
Norton records frogs then calling August 15-16, 1914. Possibly June 24-
August 16 represents the range of the mating period. In general the bulk
of mating must come from June 2 8- July 30. In 19 13 the choruses at Otter
Lake, Ontario, were from July 7-15. This year (1923) at Onekio, N. Y.
they were not mating June 14. On June 28 at Adirondack Lodge, Hart
Lake, Messrs. S. C. Bishop and C. K. Sibley found them in chorus and the
males very impulsive but no eggs present. About July 7 Mr. C. W. Leister
also heard them at Hart Lake as did Mr. and Mrs. G. B. Upton indepen-
dently. On July 14 I found the species mating and breeding at Hellgate
ponds, Onekio, N. Y. My friend Dr. S. C. Bishop at Hart Lake also found
them breeding there July 14. About the same time July 8, 1923, Dr. S. E.
Simpson at Otter Lake, Dorset, Ontario, reports by letter that he had recently
found them breeding. It would, therefore, appear that July 7-15 is an almost
certain mating date for this species if three of us independently find the eggs
July 8-14.

This species usually is most active at night. On July 14 I saw about
9:30 a.m. six or eight males spread out on the surface around a mass of
sedges. They were vigorously croaking and from time to time rushing at
each other, the rushee thinking the other to be a female. They lie spread out
with hind legs on the surface. On July 2, Messrs. Bishop and Sibley took
on Clear Lake "24 specimens of which 23 were males and one juvenal female.
These males were swimming and floating in open water 3 or 4 rods from shore
or resting on pads of the yellow cow lilies. " On July 13 and 14 we have the
following notes from Dr. S. C. Bishop. On July 13 he "took two mature
females; one specimen with eggs extruding was resting at the surface of the
water with no male in her vicinity; the other female was at the surface and
near the shore. On July 14 placed at 9:00 a.m. a male with first female
(extruding eggs) in a pan and kept dark. By 9:30 they were mated and so
remained until 2:00 p.m. when the female was found to be dead. The male
grasped the female with the first two fingers above and in front of the brach-
ium and the other two fingers behind the brachium. Later the male had
one arm wholly ahead of the female."

4o6 Frogs of the Okefinokee Swamp

The ripe females seem to be very rare and the males in the lake greatly
predominate. Immature females are more common near shore. Dr. Bishop,
as do I, believes that the females stay near shore in hiding until ripe, then
swim out toward males which rush for them or to meet them. Doubtless
then the pair sink to the bottom or soon the female seizes a stem for ovulation.
I believe the embrace is normally pectoral because of the cross embrace with
R. clamitans and the swollen thumb.

Mr. Garnier's notes on mating are "On the 24th of June I collected a
number of R. septentrionalis and placed them in a large white earthen vase.
They remained quiet for a time and I put in some chips and a quantity of
Ranunculus. Next morning three couples were paired and lying at the
bottom of the vase, and secreted among the Ranunculus. One pair were
on the surface, but the female had been injured. It thus seems they accouple
in the night, and immediately sink and hide. Occasionally there was a trivial
chant amour from the last pair, evidently so given, but the others were
mute. . . . The male seizes the female by the lower portion of the axilla, near
the upper third of the dorsal vertebrae, but not by the lumbar regions. At
this time the tinting on the chin and throat was a fine gamboge-yellow, and
was deeper toned in some specimens than others, but not particularly more
in the males than in the females. In both sexes it was equally beautiful. I
could not help being struck by the extreme stillness of the pairs in coitu
among the Ranunculus. Nothing seemed to induce them to move in any
manner. They were at rest. I carefully examined since on all opportunities,
and searched the streams and pools to find some in coitu, if possible, to observe
them in their natural embrace but as yet without success. . . .'

"Thus it may be justly inferred that after the female is grasped the pair
sink to the bottom and conceal themselves from view, and that they bury
themselves in the mud or seek the covering of water plants, after the manner
of those in the earthen vase. It is likely some prompting of nature that thus
makes them bury themselves from sight to protect themselves from enemies
that could, at that time, make them an easy prey, and in security perform
their process of fecundation. I kept my specimens referred to for over a
fortnight, but no spawn was deposited."

Amplexation, {Normal, abnormal). We have no data on the actual
mating. The very secretive nature of this animal's life, at the non-breeding
period, its attachment to cover in the day unless it be overcast would lead
one to expect it to mate at night. Most surely its amplexation must be
pectoral as its enlarged thumb would indicate. The duration of mating
must be very brief and the concourses of this specialized species shorter than
with most species.

Cross embraces with other species were not provided for. On July 14
we placed several males with two green frogs in a dark fish can. When we
reached home 12 hours later we found one male R. septentrionalis embracing
a female R. clamitans with arms dug into the axils. Its arms could not span
farther i.e., reach around on to the breast.

Rana septentrionalis 407

OVULATION

Habitat. The normal habitat for the non-ovulation period is retained for
ovulation. We clearly beHeved the eggs would be found in connection with
water Hlies, hence our choice of habitat for the mink frog in "Frogs: Their
Natural History and Utihzation" U. S. Bureau of Fisheries Report 1920,
App. Plate VIII, p. 17. Our experience at Onekio, N. Y., Hellgate Pond,
July 13 and 14 tended at first to lead us to believe Garnier's observed habitat
the preferred one but the records of S. C. Bishop and S. E. R. Simpson to
be related later prove the water lily habitat as best. Garnier (pp. 950)
writes: "Today is the 30th of July. . . . There is plenty of spawn in the
streams." On July 14, 1923, Mrs. Wright and I found plenty of frogs below
a beaver dam where the outlet of a pond flowed. Just below the dam was a
large area in which were the tadpoles of Rana clamitans and Rana septen-
trionalis, redbelhed dace {Chrosomus erythr eg aster), common shiner {Luxilus
cornutus), horned dace {Semotilus atromaculatus) , fall fish (Leucosomus corpo-
ralis), cutHps (Exoglossum maxillingua) and blacknosed dace {Rkinichthijs
atronasus). Below this pool came an area with cobblestones with water
between and under, evidently usually covered with water and normally swift.
The cobblestone area was succeeded by a broad deep area of water 21/2 feet,
brown in color, more or less clear and with sediment in the bottom which
might be easily stirred up. About this area are granite boulders amongst
which are regal's fern, deciduous holly, Mt. holly, red maple, meadow sweet,
toothed haw, tall meadow rue, Mt. maple, clematis and polypody. Here we
had the bullfrog, greenfrog and mink frogs in areas. The bullfrogs were call-
ing above the dam in the pond proper. In this broad area below the dam
on the west side was an open sunlight edge of grasses, sedges, Joe Pye weed,
and monkey flowers. Here beneath the shades of Myrica Gale bushes were
fresh masses of greenfrog eggs. On the east side near a boulder and fringe of
overhanging alders {Alnus incanus) was a group of six or eight male mink
frogs floating among some scattered, broad-leaved sedges in deep water.
Farther under the alders and all along the stream under the alders at their
bases were R. septentrionalis males. Sometimes where a mass of driftwood
might be we would find five or six gathered or they would be floating on royal
fern leaves which were on the surface. Just beneath the five or six males first
discovered we found in the sedges a mass 21/2 inches below the surface.
It was so covered with sediment we thought it farther along in development
than it proved to be. The tips of the sedges were floating in the water. We
photographed the mass in place, picked the sedge on which it was attached
and returned fifteen minutes later and searched the whole spot over again.
Here directly beneath the first mass some 3 or 4 inches was a second mass.
How we missed it is hard to understand except that the sediment so obscured
it. When we at last saw it, it looked like a brown stone encircling a sedge,
an impossibility. The mass reminded me very much of R. virgatipes eggs
of the Okefinokee Swamp or R. palustris eggs not so brown and yellow.
The first mass was a plinth 2 or 2 1/2x1 1/2 inches. The second one was no

4o8 Frogs of the Okefinokee Swamp

more than 2x2x1 inches. This species doubtless Hke others lays many
complements in special areas.

Mr. S. C. Bishop on July 13, 1923, at Adirondack Lodge "found one small
lot of eggs (12) on the sandy bottom of the lake in about 8 inches of water.
They were being eaten by a newt when found. Either the newt may have
carried them there or they may have been a few laid by a female which laid
a few before the main ovulation. On July 13 p. m., day mostly bright
and windy. Found three egg masses of the mink frog {Rana septentrionalis
Baird) fastened to the stems of the yellow cow lillies. They were attached 8,
12 and 18 inches below the surface in plinth masses from 4 to 5 1/2 inches in
largest diameter. The lilies are 4-6 rods from the shore in 6-8 feet of water.
Lost one egg mass. The egg mass is slightly heavier than the water and sinks
slowly when detached. July 14. Found 2 egg masses. This date ist — about
16 to 18 inches below the surface. The embryos already hatching. 2nd lot.
2 feet below surface, freshly laid, collected and photographed. Newts very
common in the lake are the enemies of the eggs and embryos. In mass of
hatching young one newt was found in middle of the mass. In attempting
to photograph eggs in lake newts came and attempted to crawl into the mass.
During the day many males left the open lake to hide along the shore. Eggs
are apparently laid at night — at least we could find no females in act of laying
during the day or early night when using the flashlight nor in the early
morn at 5 130 a.m.

"July 15, a.m. Dark and rainy. Three egg masses found. One lot 4 1/2-
5 feet below the surface; the 2nd with embryos ready to leave egg mass 3
feet below the surface was 6 inches in greatest diameter, and the jelly ready
to disintegrate; the third mass 18-20 inches below the surface. July 15 p.m.
Two egg masses one about 4 feet below surface in 7 feet of water was 6 inches
in long diameter and fastened to two stems of yellow lily (Nymphaea varie-
gata). The mass (well developed embryos) was longer than wide with the
supporting stems running through one side; 2nd egg mass smaller about
2x4 inches and 3 feet below the surface."

I have long felt they laid far below the surface on water lilies and Mr.
Bishop told me how he proceeded. At first he found eggs by casual obser-
vation from the boat. Then later he had some one row him about and he
laid on his stomach in the prow and in this water discovered masses 4-5 feet
below the surface. On July 16 I received the following in a letter dated
July 8 and written by S. E. R. Simpson at Dorset, Ontario:

"I believe I have found the mink frog eggs. They are in a long brown
ribbon-like mass rounded. The eggs are very small and oval shaped, not
round. They were attached to the stems of weeds either floating at the
surface or else half way between the bottom and the top. I found them at
Peat Lake all along the shore among the lily pads. If I have any luck I should
be able to get some tadpoles. There are lots of them and we got about 18
bunches." In this same place, I collected adult frogs in 19 12 and in 19 19
in the middle and last of August Mr. Simpson took sixty or more with a net
from the lily pads.

Rana septentrionalis . 409

Period. Previous to 1923 we knew nothing positive of the eggs except
Garnier's records. Several of my friends volunteered to help in the search
that summer, and at least three of us independently found the species breed-
ing in different places. Mr. S. C. Bishop had kindly invited me to join him
at Hart Lake July 13 and 14. I could not make the proper plans and he
went to Hart Lake and Mrs. Wright and I to the Fulton Chain of lakes,
Adirondacks. We found only two masses at Onekio, N. Y., the species just
beginning while Mr. Bishop reported the species well launched in Hart Lake,
Adirondack Lodge, near North Elba and Mt. Marcy. On July 16 I saw a
letter from Mr. S. E. R. Simpson who at Dorset (Otter Lake), Ontario,
reported on July 8 that he believed he had found the mink frog eggs. These
three independent discoveries July 8-14 would seem to indicate the first
of July as a part of the crestal period of ovulation. Mr. Garnier's records
June 24 and July 30, 1883, are yet the extreme dates of ovulation. We,
therefore, have from June 24 to July 30, possibly August 16 as the egg period
with the bulk of eggs laid in July. Most of the U. S. National Museum
accessions are for the month of July when this inconspicuous species is
croaking and most in evidence. U. S. National Museum No. 36097 is a
female 67 mm. and ripe. It was taken July 9, 1903, at Cross Lake, Aristook
Co., Me., by Mr, W. C. Kendall. On the same day of the month, July 9,
1900, G. S. Miller Jr., secured a ripe female 61 mm. (U.S.N.M. No. 28355)
at Peterboro, N. Y. These help to reinforce the impression that July is its
principal breeding month.

Egg-laying process. Doubtless the bulk of egg laying is at night. The
males await the coming of the females. When they approach the males rush
for them and soon they amplexate. It would seem they soon sink to the
bottom. As with other Ranas which lay plinths the laying after it has begun
probably lasts from a few to 15 or 20 minutes. As yet no one has seen the
actual process.

EGGS

We have already seen that the mass is a plinth with its greatest diameter
rarely as large as 6 inches, although 3-5 inches are the more usual dimensions.
In breeding habits the species reminds me of Rana virgatipes, each with
plinths except that R. septentrionalis has an inner envelope around each egg,
or of R. palustris which has a globular mass of eggs more brown and yellow.
R. palustris has the more brownish and more yellowish eggs; R. virgatipes
is next and R. septentrionalis eggs have a slight brownish and yellowish cast
at times. In the two masses we found at Onekio the fresher bunch had almost
black above and cartridge buff below. Later the lower color was almost
white. The older bunch had the eggs seafoam yellow or light chalcedony
yellow below and a dark brown above. Mr. Bishop held that "the upper
half in freshly laid eggs is black; the lower half creamy white, division of
color being about equal." Mr. S. E. R. Simpson speaks of them as a brown
mass — whether it is from the sediment in its general effect on the upper poles
I do not know.

4IO . Frogs of the Okefinokee Swamp

Some of the eggs taken July 14, 1923, at Onekio, N. Y, measured from
1. 3-1. 6 mm. in vitellus, average 1.4, mean 1.4; 2.4-3.0 mm. for the inner
envelope, average 2.7 mm.; and 5.6-6.6 mm., average 6.3 mm. for outer
envelope. As development proceeds the outer envelope may exceed 7.0 mm.
Mr. Bishop made a field measurement and examination. He writes "The
individual eggs have 3 envelopes visible with a hand lens: ist closely sur-
rounds the egg; 2nd about 3 mm. diameter; last 3rd 3/8 inch." The first
is the vitelHne membrane and the 3/8 inch envelope is between 8 and 9 mm.
It is possible some of the older eggs have the envelopes enlarged from 6-7
mm. to even 8 or 9 mm.

HATCHING PERIOD

The two egg masses (quite fresh) taken July 14, 1923 at Onekio hatched
July 18 and 19 or about 5 or 6 days after deposition. They were in shallow
pans with water between 60° and 70°. Doubtless in nature the hatching would
not be more rapid particularly with the deeply laid masses. On July 2, Mr.
Bishop and his associates could find no eggs at Clear Lake, Adirondack Lodge.
On July 13-15 he found several masses about at the hatching period. Pre-
sumably then II to 13 days is a maximum for this mid-summer form if the
eggs be deep in the water.

MATURE TADPOLE

Color description from life (July 11, 1923.) General coloration is citrine
or yellowish olive to dark olive or olive in specimens where hind legs begin
to develop well. On the back are small scattered dark spots more or less
uniform. When a tadpole has hind legs well developed (forelegs not out)
back becomes bluish black and the hind legs stand out by color contrast by
being a prominent citrine, buffy citrine, dull citrine or olive citrine. When
it reaches the four legged stage the spots of the posterior back are quite well
outlined. The mental region is grayish olive and more or less clouded. The
pectoral region has a little of greenish color. Where the sides join the belly
the body is mottled. Belly straw yellow, colonial buff or deep colonial buff.

Tail. On the lower crest along the edge are many roundish cartridge
buff or pinkish buff spots. On the base of the musculature of the tail these
collect as pinkish cinnamon spots suggesting somewhat the light areas in a
similar place on Rana grylio tadpoles. On the rim of the dorsal crest some
of the cartridge buff spots are almost whitish. There are very few black
specks on the tail in mature tadpoles. A little later the crests have prominent
black spots on posterior half of the tail. In two-legged tadpoles a black
blotch or blotches appear with pinkish cinnamon spots at the base of the
tail and these are very prominent.

Iris black and pinkish cinnamon.

Some tadpoles one month old had on the dorsum three pairs of black or
dark spots; one on each nostril; one on or near each eye; and one on each side
of the middle of the back. The dark of each eye connected with the back
spot by an arc or semicircle of light color. This is the general appearance
without a lens.

Ra7ia septentrionalis 411

General appearance. Tadpole large (99 mm.) of the R. grylio or R.
clamitans type, but without black gular area of R. grylio. Venter heavily
pigmented with white so intestine does not show through in life or preserved
specimens. Tail elongate, tip acute. Dorsal crest not as wide as muscula-
ture, not extending forward on body much beyond the level of the buds of
the hind legs. Spiracle sinistral, directed backward and obliquely upwards,
just visible from dorsal aspect. Spiracular opening just touches lateral
axis as in R. grylio. Eye just above or rarely on lateral axis. Anus dextral
on level with lower edge of ventral crest. Muciferous crypts distinct. A
short dorsal row of 8 to 10 pores on either side of middle hne of back from
the dorsal crest forward; from above the middle line of insertion of tail
musculature on body to back of eye is a distinct line of pores. Just before
it approaches the eye area it sends obliquely upward and backwards toward
dorsum a short line in the direction of the end of the first row described.
When this main line reaches area just back of the eye, a blind short line starts
transversely across occiput but is very short, another continues as supraor-
bital line to above nostril, another behind eye and under mouth as infraor-
bital to nostril. The portion behind the eye sends off an upward and back-
ward loop which immediately swings downward parallel with the infraorbital
for a short distance, then it descends directed downward to level of upper jaw
to be and turns at right angles forward toward the mouth. Another lateral
row forms a loop around the tip of the spiracle.

Mouth ports: Teeth 2/3. Edge of upper labium fringed with teeth,
the fringe about the length of the horny beak, or somewhat longer. In
either corner, beneath this fringe is a short row of teeth one sixth to two fif-
teenths the length of the upper fringe. The outer end of the lateral row does
not reach beyond the edge of the upper fringe. The median space between
these two rows is three and one half to four and one half times the length of
either row. From above the end of the first upper labial row down to the end
of the third row are two or three irregular rows of papillae inside the outer
row of papillae. These run to the end of the third row and in one case where
the third row is lost it is replaced by papillae inside the usual single row
across lower labium. Third lower row therefore about equal to the single
row of papillae on lower labium, much less than horny beak, and 1/3 shorter
than the first row of lower labial teeth. The first and second rows are about
equal to horny beak. The first one often subdivided. In general it is much
like R. grylio and one adult tadpole has a black line of spots on dorsal crest
much hke gi^ylio tadpoles.

Measurements. Length of body (24-33 mm.) in tail (45-67 mm.) 1.66-
2.1, average 1.86. Width (13.5-23.0) of body in its own length 1.3-2. i,
average 1.56. Depth (11-19.5 mm.) of body 1.14-1.33 in body width, aver-
age 1.23. Depth of body 1.64-2. 16 in body length, average 1.85. Depth
(12. 0-19.0 mm.) of tail in length of tail 3.2-4.7, average 3.87. Depth of tail
.85-1.1 in body depth, average 1.97. Muscular part (6.5-13.0 mm.) 1.3-1.9
in depth of tail, average 1.48. Spiracle 1.2-1.3 nearer base of hind legs or
vent region (9-18 mm.) than the tip of the snout (16.0 -22.5 mm.), average

412 Frogs of the Okejinokee Swamp

1.37. Spiracle i. 25-1. 6 nearer eye (6.5-12.5 mm.) than base of hind legs or
vent, average 1.45. Eye 1.-1.4 nearer tip of snout (8-1 1 mm.) than spiracle
(8.0-12.5 mm.), average 1.14. Nostril 1.0-1.6 nearer eye (4.0-5.5 mm.)
than snout (4.0-6.5 mm.), average 1.16. Mouth (4.5-6.5 mm.), usually
1. 0-1.3 larger than internasal space (4.0-6.0 mm.), average i.ii. Mouth
contained in interorbital distance (6.5-9 mm.) i. 3-1. 75 times, average 1.55.
Internasal space contained in interorbital space 1.4-1.9, average 1.7.
The dimensions of the largest tadpole are:

mm.

mm.

Total length

99

Spiracle to vent

18

Body length

32

Spiracle to eye

12

Body depth

19-5

Eye to snout

II

Body width

23.0

Eye to nostril

5-5

Tail length

67 .0

Nostril to snout

6.0

Tail depth

18.0

Mouth

6.0

Musculature

of tail

13.0

Interorbital distance

9.0

Spiracle to snout

22.5

Internasal distance

50

LARVAL PERIOD

Garnier in his point number 4 held that the tadpoles of R. catesbeiana,
R. clamitans and R. septentrionalis require two years to mature. In another
place he wrote: "Frog now spawned (July or August) cannot be completed
this season, as there are plenty of tadpoles in October and November of
R. catesbeiana, septentrionalis and clamitans. They are seen, all of them,
without limbs in spring, and at the present moment they are all three being
perfected and assuming the imago, or perfected form. Thus, it requires two
years to perfect this little frog."

Inasmuch as R. clamitans may transform from 365-400 days from egg
deposition doubtless Garnier is in error in believing two years necessary for
transformation in R. septentrionalis whose tadpoles transform at 29-38 mm.
or average 33 mm. R. clamitans transforms at a similar size 28-38 mm.,
average 32 mm., while R. catesbeiana transforms 43-59 mm. or 53 mm. aver-
age, the latter size, practically a one year old R. septentrionalis beyond trans-
formation. If some of the R. septentrionalis at 41-44 mm. with stumps are
not tardy in losing their tails after transformation possibly R. septentrionalis
occupies an intermediate position between R. clamitans and R. catesbeiana.
I prefer to believe one year or slightly more the larval period, yet two years in
some cases may be required.

TRANSFORMATION

This may come from June to August or September. Garnier (p. 951) "On
the 24th ult. (June, 1883) obtained several tadpoles, one a nearly perfect frog
with only a small fragment of a tail to be absorbed; several had both legs and
arms and others the hind legs with the arms quite ready to make their appear-
ance and the skin confining them at the shoulders, transparent." On July
i6th (Copeia, 1915, p. 45) Wright captured at Dorset, Ontario, "one or two
frogs with the stump of the tail remaining. These were about 38 mm. {i}/2

Rana septentrionalis 413

inches from snout to vent, or little below the two largest specimens which
were taken (49 mm. or 2 inches). The other specimens which were not lost
measured 47, 42, 42, 43, 40, 40 mm. respectively." At the time I did not
sense it but these eight were doubtless little past transformation.

On August 8-9, 191 7, Pope (Copeia, Dec. 31, 19 18, p. 96) found tadpoles
transforming and transformed at Tim Pond, Eustis, Franklin Co. Me.
"Most of the tadpoles had completed their transformation before this date
but I collected a single one with large hind legs, measuring i 15/16 inches
including the tail. Most of the frogs found had evidently completed their
change this summer and some retained the stump of a tail. They averaged
13/4 inches long, snout to vent. The adults of which I collected four, were
nearly twice this size 2 15/16 inches."

In a group of 60 or more specimens taken the middle and last of August,
1 9 19, Mr. S. E. R. Simpson has three just transformed at 32, 35 and 36 mm.
This summer (1923) from tadpoles from Onekio, I have two transformed on
July 2, two at 30 and 34 mm., and one July 14 at 29 mm. We, therefore,
have transformation from June 24 (Garnier) to the last of August (Simpson)
at from 29-38 mm. or possibly to 40-49 mm. I believe that sometimes after
real transformation they retain the tail for a considerable period. The bulk
of transformation doubtless comes in June and July.

Subsequent notes on some of the U. S. National Museum material are as
follows :

(a) Mill Creek, Sackett's Harbor, N. Y., Evermann & Clark, July 2,
1894.

No. 39747. Transforming, 4 legs. 34 mm. in body; tail stub 29.
Dark band more or less down middle of tail musculature but band
broken up.

No. 39748. Transforming, 4 legs. 29 mm. in body, tail stub 27
mm; tail musculature with broken band.

(b) Peterboro, N. Y. Mrs. E. P. Miller— 1900.

No. 28455. 37 nim. Not long past transformation.
No. 28456. 36.5 mm.

(c) Peterboro, N. Y. Mr. G. S. Miller, Jr., July 26, 1900.
Just transformed.

Just past transformation.
Just past transformation.
Just past transformation.

Just past transformation.
Just transformed.
Just past transformation.
Just past transformation.

(d) ist Connecticut Lake, N. H. Bur. Fisheries, July 21, 1904.
Still signs of transformation.

Apron on back from transformation.

(e) Newport, Clyde R., Vt. Evermann & Bean, July 23, 1894.
Just transformed.

No. 28338.

36 mm.

" 28339.

38.5 mm.

" 28340.

38 mm.

" 28341.

40 mm.

" 28342.

42.5 mm.

" 28344.

40 mm.

" 28345.

41 mm.

" 28346.

39 mm.

ist Connecticut Lake,

No. 36465.

40 mm.

" 36466.

38.5 mm.

Newport, Clyde R., Vt

No. 39814-

31.5 mm.

414 Frogs of the Okefinokee Swamp

(f) Lucknow, Ont. J. H, Garnier, July i, 1883.
No. 13605. One 30 mm., tail stub small.

One 31.5 mm., tail stub 46, dark mottling along middle

of musculature.
One 30 mm., transformed, slight stub.
All the above U. S. National Museum material apparently came from
July 2- July 26 and ranges from 29-40 in transformation size.

GROWTH

If we presume 29-40 inches to be the normal transformation size and the
bulk of transformation to be the last of June and through July, possibly
some specimens (average 44 mm.) taken the middle and last of August, 19 19,
at Peat Lake, Dorset, Ontario, may give us some clue to growth. The ma-
terial sorted into three or four groups: the newly transformed, 3 specimens,
32, 35, 36 mm.; i specimen 41-47 mm., average 44 mm.; 27 specimens, 50
mm. 9-58 mm. cf, average 53 mm., and i, 63 mm. 9. It would seem that
they transform in July at 29-38 and by the middle and last of August of the
following year they might reach 41-47 mm. in length. One year later they
may attain 51-58 or possibly 63, though the latter measurement may be a
three year old group. Cope measures 2 specimens 1.92, 49 mm., and 2.25
(58 mm.) inches respectively in body length. Pope (Copeia, Dec. 31, 1918,
p. 96) records four 2 15/16 inches (75 mm.).

From Onekio, N. Y., June 10-14, 1923, we took three groups: One of 29,
30, 33 mm. respectively just transformed; the second of 42, 48 9 (?) mm.
respectively; and the third group of 50 9 and 57cf mm. In the same place
one month later July 13, 14, 1923, we took three or four groups: 33, 34, 35,
37, 37, 38, 38, 39 mm. near transformation; 41 and 48 mm. sex undeter-
minable externally: 52 9 ?, 52.5 cf , 53 cf , 54 cf , 54 cf , 56 cf mm.; a four
group possibly 59 cf , and 64 9 mm. We then have 29-39 mm., 41-48 mm.,
50-57 mm., 59-'63 9 mm., revealed as four groups for Onekio, N. Y.

From Hart Lake or Clear Lake, Adirondack Lodge, Adirondacks, N. Y.
June 28, 1923) Drs. S. C. Bishop and C. K. Sibley secured the following
six specimens: one 53 mm.; the rest 65, 69, 70, 70, 71 mm. respectively. On
July 14, 1923, from same place we have one 51 cf mm., and two 60 9 and
63 9 mm. The 50-53 mm. specimens and 60-7 1 mm. specimens seem to reveal
two groups.

Under (Transformation) it will be revealed that in the U. S. National
Museum are 7 or 8 series in which are transformed or past transforming
frogs. They range from 29-34 mm. (Sackett's Harbor, N. Y.) to 36-40 or
possibly 42.5 mm. (Peterboro, N. Y., G. S. Miller, Jr.). When Mr. Miller
took these he also secured a 58 9 mm. specimen or when Mrs. E. P. Miller
secured 36.5 and 37 mm. specimens she also secured 41, 41, 41, 48 9, 47 9
mm. specimens. One other lot secured here on July 9, 1900, is 51 cf mm,
56 cf mm., 57 cf mm.; and 61 9 mm. and 66 9 mm. This Peterboro ma-
terial shapes as follows: 36-40 mm., possibly 42.5 mm. for transformation;
47 9 or 48 9-58 9 mm.; 61 9-66 9 mm.

RaTia septentrionalis 415

The whole U. S. National Museum series shapes thus 29-40 or possibly
42.5 mm. transformation, average 37 mm.; 41 or 42-47 or 48 mm., average
43.5 mm. first year olds; 48.5-58 or 58.5 mm., average 53.5 mm. two year
olds; 60-72 mm., average 64 mm. three year olds. This material and ours
combined gives 29-40 mm., average 37 mm., mode 38 mm. at transformation;
40-48 or 50 mm., average 44 for ist year olds; 47 or 48 or 50 -58or 58.5 mm.,
average 53.5 mm., mode 51 mm. for two year olds; and 59 or 60 to 72 mm.,
average 63 mm., for three year olds.

FOOD

Garnier finds that (p. 945) "it preys on water beetles and similar insects,
but seems especially partial to the Julus family, having generally found it
in the stomachs examined." (P. 947) "the stomachs of many have been
examined by me, and they contained mostly Carabus, Julus, and water
insects, and on two occasions some little fish, chubs, if I remember correctly,
about an inch long. Thus their food is hke that of other frogs." In two dif-
ferent instances he found tadpoles eating dead fish, one a brook trout, the
other a chub. He saw one (tadpole) rush at, seize a large Ephemera that
came near it and swallowed it, yet it had not more than the third of its tail
absorbed."

ENEMIES

As already revealed under eggs the newt is a serious foe of the eggs and
hatching embryos. Being more common in the southern parts the water-
snake is a negligible factor in the life of this species unlike the case of R.
virgatipes. The great blue heron and other herodiones are amongst the worst
foes of the mink frog.

AFFINITIES

Garnier considered "certain peculiarities in the life history and in external
forms of these three Ranas {R. catesbeiana, R. clamaia and R. septentrionalis)
which so thoroughly agree that they may be separated into a group by them-
selves. These I shall endeavor to point out as concisely as possible:"

"i . They have no 'chant amour' or love notes in spring.

"2 . They retire early to hibernate with the first autumnal frosts.

"3 . They live in water and lie in wait for their food, but do not hunt for
it on land. They poise the body on any floating weeds, lie on the
bank or any bit of stick or log that suits their purpose.

"4. The tadpoles of R. catesbeiana and R. fontinalis require two years to
mature, and the mink frog requires the same period.

"5. Adults in all three have no lateral fold, but merely a slight raising
of the skin from the angle of the mouth, and which terminates or
shades off on the shoulder.

"6 . The foot is broader in proportion than in the rest of the family, and
the second toe is proportionately shorter, a peculiarity emphatically
distinct, and can be seen at an glance by any one who takes the trouble
of even a cursory observation, webbed to extremities.

4 1 6 Frogs of the Okefinokee Swamp

"y . When captured they sometimes utter a cry of distress quite different
from their ordinary croaking notes, and I have often seen a bullfrog
open his mouth and scream for even a minute, like a child in distress.

"8. When they give their note it is always produced by inflating the
throat pouch and suddenly expelling the air, whereas in R. halecina
there is a pouch near the angle of the jaws, on either side.

"g. They are all tinged, more or less, with yellowish-green on the chin,
which soon shades towards the throat and breast, and on the belly
is white, more or less, in many subjects most beautifully so.

There is thus an analogy in their life history, and in their external confor-
mation that at once forms them into a group by themselves, and makes a
marked section. I am not aware, however, that there is any anatomical
difference sufficient to make a genus."

In 1889 Cope (pp. 396, 397) graphically makes this species a central form
from which four lines of North American Ranas radiate. "Sometimes the
vomerine teeth in Rana temporaria pretiosa are not appreciably more pos-
terior in position than in Rana septentrionalis, in which case the species
approach each other very closely. The Rana septentrionalis violates the
characters which distinguish the R. clamata and R. caiesheiana from each
other, and would afford a complete connection between them were it not
for its inferior size; but even this point does not invariably hold good, as a
few specimens of R. clamata do not exceed it in dimensions."

"These series (four radiate lines above) are not probably genetic, as some
of the species have been most likely derived from the Old World. The R.
septentrionalis, however, may be very probably ancestral to the forms of the
Catesbeiana series, and perhaps of others."

"The variations of the Rana septentrionalis are greater than those of any
other North American species of the genus. There is, however, no coinci-
dence between them, so that they cannot be regarded as indicating subspecies.
The tympanic disc varies greatly in size, the males having it larger than the
eye, and the females smaller than the eye. In this respect the species displays
its near affinity to the R. clamata and R. catesbeiana. Some specimens have
a dorsolateral dermal glandular ridge, and others have none, such difference
being exhibited by specimens from the same locality. The spotting of the
dorsal surface varies very much. In some specimens the spots are not closely
placed; in others they leave only narrow fines of the lighter ground color
between them."

Taking all its characters together, this species occupies a position inter-
mediate "between nearly all the North American species of the genus, and
from some such form it might be supposed that all the Ranae of the northern
hemisphere have been derived." (Pp. 418, 419).

Boulenger (1920, p. 425) later comments: "I cannot agree with Cope . . .;
nor do I think him justified in adding that 'from such a form it might be
supposed that all the Rana of the northern hemisphere have been derived,'
a statement I would rather apply to Rana catesbeiana and Rana grylio, re-
garding R. septentrionalis as connecting R. catesbeiana with R. clatnitans."

Rana septentrionalis 417

There is another point of view expressed in H. H. T. Jackson's "The
Land Vertebrates of Eidgeway Bog, Wisconsin: Their Ecological Succession
and Source of Ingression. (Bull. Wise. Nat. Hist. Soc. Vol. 12, Nos. i and 2,
June 1914, pp. 17, 18). "The northern frog, Ra7ia septentrionalis, has been
considered by some herpetologists to be a subspecies of Rana clamitans;
specimens of the two forms are often morphologically very similar, but their
habits and behavior are so widely at variance that it seems improbable they
interbreed. For example, Rana septentrionalis is relatively, a sluggish frog;
during the summer months it is found along the edge of lakes and streams,
usually with the body partly submerged in the water, occasionally on land
at the water's edge; when alarmed it utters no sound, makes a feeble leap
of two or three feet into the water, settles to the bottom and hides in the mud
or under some object where it is easily captured in the hand. Contrast with
the behavior of this frog that of Rana clamitans] the latter is an active frog;
during the summer months it is almost invariably found out of the water on
a gently sloping bank; when alarmed it utters a loud startling croak, makes
a vigorous leap of five or eight feet into the water, swims some distance
under water before hiding, where it remains alert and difficult to capture.
There remains, of course, the alternative to explain this as "one species
having different mores" (Shelford, 1912a, p. 92), but one can hardly accept
this view in the present case."

Such discussions are unsatisfactory. It depends on what characters one
knows best, what he is most interested in, or what he accidentally discovers
that he uses as his criteria or canes on which he leans. What follows may
be of no consequence :

1 . Little evidence can link it with Rana sylvatica or R. cantabrigensis.
Sometimes the adults are almost free of spots like Rana sylvatica,
R. cantabrigensis, or some of the Northwestern species. Its eggs are
with two jelly envelopes and the above species have the second
indefinite. The egg mass is submerged as in R. sylvatica, or R. ca7ita-
brigensis and about of the same size. The woodfrog and northern
woodfrog choose woodland pools, etc.; R. septetitrionalis, peaty lakes,
waterlily ponds. The tadpoles are wholly unlike the woodfrog, one
transforms the year laid and the other after wintering over at least
one season. The two have deep tail crests far forward in body and
with labial tooth rows more numerous, while R. septentrionalis has
the elongate large narrower tail crests of a Rana clamitans type and
tooth rows 23. The two transform at a small size; R. septentrionalis
transforms at a variable but usually large size. There is little in
common with the woodfrogs.

2 . Rana pahistris.

The pickerel frog (Rana palustris) is one of our most poisonous
frogs with a decided odor. So also Rana septentrionalis is the "mink"
frog. Therein they might be associated if secretions be the criterion.
Each lays submerged egg masses, each has eggs quite similar in size.

4i8 Fj'ogs of the Okefinokee Swamp

envelopes and color. Each adult has about the same degree of vocal
sac development in the male.

The adults are unlike in color pattern but Rana septentrionalis
may have the upper parts with pronounced spots but not regularized.
Rana palustris tadpole may be held as much R. clamita7is-\\ke in
appearance as R. pipiens — (also sphenocephala, aesopus, areolata)
like but it transforms the same year as eggs laid. The transformation
size is smaller and it breeds in late April and May while R. septen-
trionalis belongs to the June- July season of R. damitans, R. virgatipes,
R. grylio, R. catesbeiana. But Rana caiesheiana breeding in February
1925 in San Antonio, Texas, cured the author of over emphasis on
this prop.

3 . Rana damitans.

Sometimes it has a suggestion of a dorsolateral glandular fold;
its tadpoles winter over at least a season, and the tadpoles have the
outline of R. damitans type; sometimes adults have no spots like
Rana damitans; sometimes R. damitans in Adirondack and Northern
parts are much spotted.

Usually they have no dorsolateral fold, are usually spotted, trans-
form at much larger size than R. damitans though adult is smaller:
lays submerged egg mass, not surface films. Its nearest relationship
lies doubtless with No. 4.

4 . Rana virgatipes.

(See Rana virgatipes Affinities).

In transformation size Rana septentrionaUs approaches Rana
catesbeiana or Rana grylio more than R. damitans, while R. viigatipes
goes into the R. pipiens and R. palustris class.

5 . Rana pipiens.

Rana septentrionaUs is an associate of this form in the west of its
range and may have come from this widespread species. Its egg
mass looks more like that of Rana virgatipes than R. pipiens yet
Rana septentrionaUs has two envelopes on each egg as R. pipiens but
thisis not to be overemphasized. They are in different tadpole and
transformation classes though the adults of the mink frog are smaller
than R. pipiens However they may approach. Witness A. C.
Weed's description of Rana kandiyohi from Minnesota where Rana
septentrionaUs and R. pipiens are. He writes: "Typical specimens
of this species show a color pattern which suggests a blending of
Rana pipiens and Rana septenirionaUs. It is as though the black
spots of Rana pipiens had been superposed on the mottled color of
septentrionaUs. ." (Weed 1922, p. 109).

BIBLIOGRAPHY
1920 Boulenger, G. A. A Monograph of the American Frogs of the Genus Rana. Proc.

Am. Acad. Arts and Sciences, Vol. 55, No. 9, August 1920, pp. 423-425.
1889 Cope, E. D. The Batrachia of North America, U. S. N. M. Bull. No. 34, Wash.

1889, pp. 416-419.

Rana syhenocephala 419

1906 Dickerson, M. C. The Frog Book, 1906, pp. 224-225.

1883 Gamier, J. H. The Mink or Hoosier Frog. The American Naturahst, XVII, Sept.
1883, No. 9, pp. 945-954-

19 14 Jackson, H. H. T. The Land Vertebrates of Ridgeway, Wisconsin: Their Ecological
Succession and Source of Ingression. Bull. Wise. Nat. Hist. Soc, Vol. 12, Nos. i
and 2, June 19 14, pp. 17, 18.

1898 Mearns, E. A. Bull. Am. Mus. Nat. Hist., Vol. X, p. 325.

1915 Pope, P. H. The Distribution of the Northern Frog, Rana septentrionalis Baird in
Maine. Copeia, No. 16, March 15, 1915, pp. i, 2.

1918 . A New Record for Raria septentrionalis Baird. Copeia, No. 64, Dec. 31,

1918, pp. 96, 97.
1922 Weed, A. C. New Frogs from Minnesota. Proc. Biol. Soc. Wash., Oct. 17, 1922,

Vol. 35, pp. 107-110.
1915 Wright, A. H. The Mink Frog, Rana septentrionalis Baird, in Ontario. Copeia

No. 23, October 15, 1915, pp. 46-48.
1920 Wright, A. H. The Frog. U. S. Dep't Commerce, Bureau of Fisheries, Doc. 888,

1920, pp. II, 16, 34, 35.

Rana sphenocephala (Cope)

(PI. Ill, Fig. 7; V, Fig. 12; IX, Fig. 3; XI, Figs. 9, 10; XIII, Fig. 2; XVI, Figs, 7, 8; XVII;
XL; XLI, Text Figs. 1,21)

COMMON NAMES

Southern Leopard Frog. Southern Meadow Frog. "Spring Frog." Spotted
Frog. Water Frog. Shad Frog.

RANGE

Check list. "Range: Southeastern States." (Stejneger & Barbour 1923,

P- 37)-

Supplementary records. Deckert (1914, Nos. 3 & s) gives two meadow
frogs at Jacksonville, Florida, each abundant. Whether it be they were
puzzling or the two were there or the distinction between the two species be
poor we cannot understand. Those just north of Jacksonville, Florida, in
Okefinokee Swamp are of the R. sphenocephala form. In 1922 the same author
(p. 88) finds only Rana sphenocephala in Dade Co., Florida. Of these he
"saw numbers along the shore of the brackish canal on the right of the
Ingraham Highway, between Royal Palm Hammock and Cape Sable. Loding
(1922, p. 20) records it in three counties in Alabama. Viosca, Jr., in 1918
(p. 160) records only R. sphenocephala but in recent years I believe he has
found both R. pipiens and R. sphenocephala as has Ortenberger (1927, pp. 46,
47) in the panhandle of Oklahoma. Edith R. Force (1925, p. 25) recorded it
from Okmulgee Co., Oklahoma. Philip H. Pope (1919, p. 97) carried it west-
ward into Texas as far as Houston, Texas. Ortenberger (1921, p. 75) found
each species R. pipiens and R. sphenocephala in Indiana. The recorded range
northward along the Atlantic Coast is to North Carohna where (G. S. Myers,
1914, p. 60) and Brimley & Mabie (1925, p. 15) record it. In 1923 we saw
meadow frogs at Lakehurst, N. J. and in southern New Jersey strikingly
like Rana sphenocephala.

Local Okefinokee records. In 1 9 1 2 we found it on the outskirts of the swamp
as well as in the swamp. We secured mainly tadpoles and young in 1912.
In 192 1 we recorded it on Billy's Island, Chesser's Island, Suwannee Canal,

420 Frogs of the Okefinokee Swamp

Honey Id. Prairie, mainland east of Chesser's Island. In 1922 we took it
along St. Mary's River, Chesser Island and mainland between the swamp and
St. Mary's River.

GENERAL APPEARANCE

We are provisionally keeping this form as a separate species. If this were
a taxonomic paper we would make more measurements and studies to solve it.
The southern meadow frog is an alert, active, long-legged and long-snouted
spotted frog.

Miss Dickerson chose the long head, circular white tympanic spot and
dorsal spots not outlined by white as distinctive of this species in contrast
with Rana jnqnens, the northern meadow frog. Boulenger, G. A. (1920, pp.
438, 439) points out that these characters are not constant and does not
recognize R. sphe?iocephala as separate. Of it Miss Dickerson writes (1906,
p. 187): '"The head is long and pointed, with the eyes set far back. This
characteristic and the usual length of the hind legs distinguish this frog at
once from Rana pipiens, the Common Leopard Frog. It is peculiar also in
possessing a circular white spot at the center of the ear. This spot is never
lacking, no matter what the coloration of the frog may be at the time. Ra7ia
pipiens has sometimes a light blotch at or near the centre of the ear, but never
this clean-cut circle of white."

COLORATION OF SPIRIT SPECIMENS (19 1 2)
In 191 2 we made these distinctions between R. pipiens and R. spheno-
cephala on the basis of coloration of spirit specimens. On the back the color
may be deep olive to green or purplish brown. On the inner portion of the
upper eyelid the blackish brown spot is smaller than in Raiia pipiens and not
so constant. Two or three of the specimens have these two spots absent.
In most R. pipiens there is a median spot ahead of these two orbital spots
and in all these R. sphenocephala this is absent. The dark line from snout
through the nostril to the front of the eye is possibly more constant and
prominent than in R. pipiens. Between the lateral folds are two rows of
spots more widely separated than in R. pipiens and less regularly placed.
Below the lateral folds in R. pipiens there are two or three irregular rows of
smaller spots but in R. sphenocephala there is usually one row followed by a
more or less vermiculated area, the black sometimes forming more or less
horizontal hues. On the fore-arm of R. pipiens there are two or three separate
spots on the posterior edge of the brachium while in R. sphenocephala these
are united into one broad brown band; in both there is a brown band on the
anterior side of the brachium, in R. sphenocephala averaging larger than in
R. pipiens. In both there is usually a prominent spot on the front of the
forearm and usually a band on the posterior edge. These rarely may be ab-
sent. The bars of the hind limbs are farther apart and less developed in R.
sphenocephala than in R. pipiens and the small spot below and in front of the
anterior femoral bars are more or less separate in R. pipiens but generally
unite to form a prominent antefemoral dark stripe, particularly in the adults
of R. sphenocephala. This femoral band often unites with the dark vermicula-

Rana sphenocephala 421

tions or reticulations of the groin. The posterior faces of the thighs in R.
pipiens have the brown spots more or less separate and the lighter interspaces
are equally emphasized to form the reticulum but in R. sphenocephala the
brown spots become connected and surround the lighter inter-spaces, the dark
being the reticulum. The stripe from the snout under the eye to the shoulder
is yellowish, much lighter and more prominent than a similar more or less
bronzy line in R. pipiens. The same difference obtains in the color of the
lateral fold. On the upper jaw of R. pipiens there is a more or less sharply
defined dark line below the above light stripes and below the dark line a clear
white or yellowish white edge to the upper lip. In R. sphenocephala these two
areas are not sharp and the upper and lower lips are more or less dark and
light spotted. In R. pipiens the lower lip is usually with a few or no dark
spots while in i?. sphenocephala this is the regular coloration. On the throat
region and under side of the hind legs dusky fleckings much more frequently
enter in the coloration of R. sphenocephala than in R. pipiens. The under
parts are whitish or yellowish white.

In most of the specimens the clear cut distinct white spot in the middle of
the tympanum is sufficiently distinctive but in some young and adults the
character fails to separate R. sphenocephala from R. pipiens. Our specimens
have not the tibia with complete cross bars but we question its use as a con-
stant distinction because many R. pipiens are without complete bars though
this latter species inclines more towards the barred state of affairs.

If time were at hand to study as carefully 192 1 and 1922 material possibly
some of these foregoing distinctions made in 19 13 would be changed or
recognized as non-distinctive.

STRUCTURAL CHARACTERS (WRITTEN IN 191 2)
On the basis of our 191 2 specimens these characters are noted. The skin
is smooth; on either side of the back is a prominent dorso-lateral fold. Be-
tween these two folds on the back there may be no ridges at all or five or six
smaller secondary ridges or folds colored like the body and not as the dorso-
lateral folds and more or less broken in appearance. These intermediate folds
are more prominent in the young specimens. The muzzle is more acuminate
than in R. pipiens. A prominent fold below the eye to below the tympanum
in breeding males or to shoulder in others. Nostril about equidistant from the
end of the snout and eye. First finger longer than the second and about equal
to the fourth. Third finger larger than the rest. A tarsal fold more or less
present; fourth toe of R. sphenocephala longer than in R. pipiens, sometimes
longer by one or two phalanges in specimens of the same size. Three phalanges
of the 4th toe free. Inner metatarsal tubercle present. Tongue prominently
notched behind; vomerine teeth between inner choanae and in transverse
series. These characters not verified with 192 1 and 1922 material.

MEASUREMENTS

(Recent Material)
Head to angle of mouth .95 (28 mm.)-.96 (44 mm.)-i.o2 (56 mm.)-i.i3
(68 mm.)-i.i (82 mm.) in width of head; head to rear of tympanum .8-.83-

42 2 Frogs of the Okefinokee Swamp

.91-.93-.86 in width of head; head to angle of mouth 2.66-2.83-2.73-2.72-3.5
in length of body; head to rear of tympanum 2.24-2.5-2.43-2.5-2.82 in length
of body; snout .66-.88-1.0-.96-.84 in ist finger; snout .66-.5-.62-.46-.60 in
4th finger; snout .41-.55-.s7-.7-.64 in first toe; eye . 87-1. 0-.85-1. 07-1. 06 in
tympanum; eye i. 5-1. 63-1. 5-1. 85-1. 6 in snout; eye 1.0-1.4-1.5 in first finger;
tympanum 2.28-2.0-2.33-2.0-2.1 in intertympanic width; tympanum 1.7-
1. 63-1. 75-1. 73-1. 77 in snout; internasal width .85-.88-1.0-.93-1.05 in upper
eyelid width; interorbital width i. 4-1. 28-1. 3 7-1. 3-1. 3 in internasal space;
interorbital width 1.2-1.14-1.37-1.2-1.4 in upper eyelid width; interorbital
width 3.2-3.14-3.5-3.0-3.6 in intertympanic width.

Forelimb: 1.86-2. 0-1.75-2. 08-2.1 in length (snout to vent); forelimb 2.73-
3 -3-3 -5-3 -7-3 -33 in hind limb; first finger 1.3 7 5-93-8 5-.80-1.1 in third
finger; second finger i. 8-1. 33-1. 4-1. 33-1. 29 in third finger; second finger 1.13-
1. 45-1. 7-1. 66-1. 15 in first finger; third finger i.o-i. 26-1. 44-1. 4-1. 25 in second
toe; fourth finger 1.375-1. 66-1. 35-1. 66-1. 5 in third finger; fourth finger .63-
1.2- 92-1. 5-1. 07 in first toe; internasal width 1.14-1.7-1.9-1.94-1.68 in first
finger; i.o-i. 22-1. 08-1. 16-1. 46 in second finger; i. 55-1. 66-1. 8-1. 55-1. 85 in
third finger; 1.1-1.0-1.17-.92-1.22 in fourth finger.

Hindlimh: length of body i. 46-1. 7-2.0-1. 8-1. 58 in hind limb; tibia 1.86-
1.7-1.55-1.45-1.82 in length; 2.73-2.84-3.1-3.02-2.88 in hind limb; 1.0-.84-
.88-.81-.86 in fore hmb; 1.33-.88-.91-.93-.91 in hind foot; first toe 2.2-1.9-
2.16-1.55-1.78 in second toe; 3.0-2.8-2.75-2.27-2.4 in third toe; 4.4-4.3-4.75-
3.77-4.0 in fourth toe; 2.2-3.7-3.16-2.77-2.5 in fifth toe; second toe 1.36-
1. 47-1. 3-1. 47-1. 4 in third toe; 2.0-2.26-2,2-2.43-2.23 in fourth toe; i. 27-1. 9-
1. 44-1. 70-1. 40 in fifth toe; third toe i. 44-1. 53-1. 7-1. 7-1. 5 in fourth toe;
.93-1.32-1.15-1. 2-1.0 in fifth toe; fourth toe 1.8-1.06-1.12-1.1-1.3 in hind
foot; 1.36-1.2-1.26-1.17-1.3 in tibia; i. 36-1. 02-1. 1-.95-1. 16 in forelimb;
fifth toe 1. 55-1. 16-1. 5-1. 4-1. 6 in fourth toe; internasal width .7-1.1-1.1-1.36-
1.3 in first toe; 1.7 5-2. 1-2. 3 5-2. 1-2. 3 in second toe; 2.03-3. 1-3. 0-3. 01-3. 2 in
third toe; 3. 14-4.8-5. 1-5.2-5. i in fourth toe; 2.0-4.1-3.2-3.7-3.2 in fifth toe.

HABITAT

In 191 2 we did not find it commonly in the lakes, runs or prairies. Its
habits accorded with those of our northern R. pipiens and we did not particu-
larly study the species. More young and tadpoles than adults were taken.
They were either in or about the small ponds and pools along the edge of the
islands, or in the cypress thickets about the islands. In 192 1 and 1922 we
made most of our records in cypress ponds or temporary pools, though
prairies, runs, Suwannee Canal, river swamps and overflowed roads and
ditches and railroad ditches also added to our localities for the species. In
the swamp it is ideally of the edge of cypress ponds and bays or small pools
and ponds at an island's edge.

Wm. Bartram (1791 pp. 278, 279) describes it thus: "The shad frog, so
called in Pennsylvania, from their appearing and croaking in the spring
season at the time the people fish for shad : these are a beautiful spotted frog,
of a slender form, five or six inches in length from the nose to the extremities;

Plate XL
Southern leopard frog (Rana sphenocephala)
Male croaking, Billy Id., Ga. June 30, 1921. Flashlight.
Eggs, stained, Chesser Id., Ga. July 13. 1922.

3. Male croaking, Billy Id., Ga. May 22. 1921. Flashlight.

4. Egg mass in camp, Billy Id., Ga. April 24, 192 1. x 1.3.
Female, Billy Id., Ga. April 29, 192 1. Ventral aspect, x 0.3.
Female, Billy Id., Ga. July 4, 192 1. Lateral aspect, x 0.3.
Male, Billy Id., Ga. May 18, 192 1. Lateral aspect, x 0.3.

*• . •• • ^^
• • • . • • •• •
• ••• ••••••.

* • • • • •

• •• • • • . • •

• • • •

•••

Rana sphenocephala 423

of a dark olive green, blotched with clouds and ringlets of a dusky color;
these are remarkable jumpers and enterprising hunters, leaving their ponds to
a great distance in search of prey. They abound in rivers, swamps and
marshes in the southern region; in the evening and sultry summer days,
particularly in times of drought, are very noisy, and at some distance one
would be almost persuaded that there were assemblies of men in serious de-
bate. These have also a sucking or clucking noise, like that which is made
by sucking in the tongue under the roof of the mouth."

FIRST APPEARANCE

In 1910, 1912, 1913, 1914, 1921, 1922 we have records of it from April to
October, December and January. In the latter months Francis Harper heard
it croaking and in December (19 13) J. Chester Bradley found both adult
males and females abroad. The months we lack records of it is more because
of absence from the swamp than from hibernation by the species. Mr. Percy
Viosca, Jr., of New Orleans, La., once told me he believed it might breed
every month in the year in Louisiana unless it be one month. Quite the same
is possible in the Okefinokee Swamp. Deckert (1914, p. 5) at Jacksonville,
Fla., finds it "everywhere from February to November." Later Deckert
(192 1, p. 21) found it breeding in December at Lemon City, Fla.

GENERAL HABITS

Some of the residents of the swamp maintain its common name of "Spring
Frog" is from its ability to leap or syring. Others hold it likes spririgy places
but usually it is called "spring frog" from its time of breeding and croaking
in the spring months. It is agile and hard to capture in water or on land.
Barbour (1920, p. 56) remarks that "Rana sphenocephala, when alarmed
while resting on a bank of a stream or pond, usually turns about and escapes
inland with leaping bounds, evidently more afraid of enemies in the water
than on the land."

On June i, 192 1, we remarked that Rana sphenocephala were hopping
around the island after breeding, also that they were becoming browner when
afield and as the season progressed.

VOICE

Of the croaking position we recorded on April 23, 192 1 that "Male Rana
sphenocephala lie on the surface when croaking or are amongst pickerel weed
stems." The rear of the body usually sprawled out (see illustrations) may be
at the surface or submerged. The head and upper back are usually emergent.
At times they may croak from the edge of a pond like R. pipiens or croak
beneath the surface or lie amongst drift wood. On May 16 we "heard several
males but they were hard to find and ceased croaking after we found them.
At 11:20 p. m. we heard Rana sphenocephala. Strange we did not hear them
when we were near the pond but distance had to sift out the calls. Over in
the north part and northeast edge of a pond were males calling amongst the
weeds. When a person approaches normally they duck or hop or swim away.

424 Frogs of the Okefinokee Sivaryip

We are having less success in photographing them than croaking males of any
other species encountered." In 1922 we were of the same opinion according to
our note of July 13. At 12.30-1:00 a. m. "We awoke and heard a great
chorus over in the area northwest of our spring. Here in grassy borders of
open spots were plenty of meadow frogs. These would stop croaking when we
approached near them. They are surely shy and begin croaking late at night."

One month later, August gth, heard two immense choruses, one at 3 :oo
a. m. and another at 4-5:00 a. m. Went to the latter congress. "Perched on
logs, sticks, and around bases of bushes were plenty of croaking males."

This species begins croaking and breeding early in the season long before
we arrived in the swamp in April. The earlier choruses come by day as well
as at night. On April 23, 192 1, it began at night at 8 or 9:00 p. m. A month
later. May i8-May 20 heard it at 8:00 p. m. On May 19 we heard them at
I :oo a. m., on June 7, at 2 :3o-2 :oo a. m., July i late in morning before dawn,
July 3 at 2:00 a. m. July 5 at 12:30-1:00 a. m., July 25 before dawn. As
the season advances the later breeders may be heard only in late morning
before dawn except for rare periods in day time before a storm passes or after
its passage.

A resume of the croaking for 192 1 and 1922 follows:

1921
April 23. Began croaking in chorus in cypress pond. Air 58°, water 60°.
Began about 8:00 p. m.
" 24. Heard several meadow frogs.
May 16. Began to croak after storm had passed.
" 18. Calling at 8:00 p. m. At times an immense chorus.
" 19. Several calling 1:00 a. m.
" 21. In Bufo pond the loudest chorus yet, 8:00 p. m. Air 70°. In several

ponds they are calling.
" 27. One heard.
June I. Hear some.
" 6. Going strong late in the night.

" 7. Croaking strongly about 2:30-3:00 a. m.

" 23. Did not hear any tonight.
" 29. Several heard in midafternoon before thunder storm. Several calling

at night.
" 30. Tonight frogs going strong.
July I. Calling in late morning before day. Few calling during day.
" 3. Calling loudly at 2:00 a. m., Honey Island. A few calling tonight

on Billy's Island.
" 5. Chorus in Long Pond 12:30 a. m.

" 6. A few croaking after midnight.

" 13. Heard one in cypress bay.
" 17. Plenty calling.

" 25. Late at night before dawn many heard.
" 29. Plenty of meadow frogs calling on Black Jack Island.
" 30. Species calling.

1922
July 13, 12:30-1:00 a. m. Great chorus northwest of camp Chesser Island.
Also a chorus in Coat Bet Pond.
" 21. Several calling in the evening.

Rana sphenocephala 425

Aug. 9. 3:00 a. m. Heard a large chorus early this morning. 4-5:00 a. m.
another chorus in Coat Bet Pond.
" II. Plenty in chorus in Coat Bet Pond 1:30 a. m. Air about 72°.
" 20. Chorus before daybreak in Coat Bet Pond.
" 22. Same.
" 29. At dusk and later a few croaking on prairie.

In the 192 1 croaking record the temperatures for the day of croaking were
from 80-97° for maxima and 52-74° for minima. Most of our camp records
for the day time range from 66-79°. Most of the large choruses came during,
preceding or subsequent to large downpours of rain. On April 23, they were
croaking in chorus when the air was 58° F. and water 60°. Inasmuch as most
choruses came at night maxima probably play the most important role of
temperatures. The average for all minima for croaking days is 65°. No
doubt that or lower temperatures prevailed when southern leopard frogs
croak. Humidity is a more important factor than temperature.

The croak itself comes from the swift inflation and deflation of the vocal
sacs (each between forelegs and angle of mouth). On April 23 we noted that
"the croak of a male sometimes is two croaks, often one or two lower croaks
or two or three. Males frequently give two or three clucking-like notes often
followed by two or three croaks. Whole performance no more than 4 or 5
seconds. On May 16 we wrote "The croak is 3, 4 or 5 croaks with two or three
clucks afterwards. The process may occupy 5 or 6 seconds." Mr. Harper
characterizes the croak as follows: "The frog may commence with a series of
guttural grunting croaks: wank-wank-wajik, ... As a whole chorus tunes
up and the clamour increases, the notes become louder and take on a more
rattling quality: wonk-wonk-wonk. They are easily heard a quarter of a
mile, . . ."

MATING

Male {From life, July 3, 1921.) Olive green or jade green or grass green on
back, pale chalcedony yellow on upper lip, pale dull green yellow costal fold;
center of tympanum chalcedony yellow. Some amber yellow or primuline
yellow on under side of fore arm or httle on the side of the groin, little on
outer side of hind legs and back of crus. Under parts white. Thumb with
black swelling. Iris: back part chartreuse yellow; upper part vinaceous buff;
inner ring amber yellow or wax yellow; rest black.

Female {From life, June 21, 1921). Top of head to back of eye cinnamon
brown to Front's brown or snuff brown to bister or sudan brown to brussels
brown. This color somewhat along inner and outer side of the costal fold, on
upper jaw from snout to front of eye somewhat on upper fore limb and some-
what amongst ground color of back. At times it makes most of the ground
color of the back. Back ground color of dorsum dark greenish olive to ivy
green. Spots of back and sides, fore arms, shoulder bar, and hind legs, black.
Those of the back with a thin hght green yellow border. Under parts white.
Throat and pectoral region dulled with blackish. Lower jaw rim marguerite
yellow or sea foam yellow with black. Same color for upper jaw below the
stripe from lower eye to arm insertion. Between the sudan brown of the side

426 Frogs of the Okefinokee Swamp

of the snout and top of the head, and in front of eye through nostril is blackish
or greenish. Two or three marguerite or sea foam yellow spots behind the
eye. Tympanum argus brown with marguerite yellow or sea foam yellow
center. Stripe from under eye over angle of the mouth and just beneath
tympanum to over arm insertion marguerite yellow or sea foam yellow.

Iris in front, below and behind pupil black with some pale vinaceous lilac
or light pinkish lilac. Rim pinard yellow. Iris above pupil maize yellow.
Very top of eye a small black area.

Structural differences. The males begin to show thumbs much enlarged
and lateral sacs well developed when 52-58 mm. in length or when presumptive
2-year olds. Our largest females are from 76-81 or 82 mm.; our largest males
from 75-78 mm.

Duration. Most mated pairs brought in laid the night of their capture
and were broken the next morning after ovulation. In a few cases where
impulsive males were brought in with unmated females they persisted in
embrace for two or three days. Observations made for R. pipiens in Ithaca
doubtless obtain for this species. The females probably do not come to the
croaking congress until ready to lay, and this may require no more than
15-30 minutes.

Night or day. From April to September it came mainly at night, in
latter part of the night, or early in the evening from April to June. Rarely
by day when a thunder storm was imminent or after it had passed.

Amplexation. — On May 16 we collected some of our first mated pairs.
These were all pectoral in embrace and we have recorded no other form of
amplexation in the species. No opportunity was given for cross embrace with
other species.

OVULATION

Habitat. In 192 1 we made these notes on places of ovulation. On April
24, we found them in a cypress pond. "Center of pond is clear (alligator
hole). Then came a circle of Pontederia in which are toad and Rana spheno-
cephala eggs. A circle of sedges; next came bushes; a thin Hne of gums; and
finally the pines." On April 28, of another pond we have these journal notes
"Frogs eggs . . . about 8 feet from edge of the pond in water 4-6 inches deep.
Two more isolated masses found each at or just below the surface and en-
circling the hzard's tail (Saururus) which is now in bloom." May 13 one of
the boys found eggs in a cypress pond. On May 22 the boys found two masses
in ditches beside the railroad ; also we found them in edge of cypress bay. In
some of the ponds just within a hammock's edge found two masses. Later
we found this form breeding in a sphagneous bog on one of the prairies.

Occasionally after severe rains they lay in very temporary places. On
June 6, 192 1, we found them in a temporary pool in the street. On July 7
we discovered four masses in an open used field just north of camp and near
a wood pile.

In 1922 we found masses in water filled pits on high oak-shaded bluffs,
beside the open road, in an open pond with saw palmetto border, but no trees.

Rana sphenocephala 427

in river swamps, ponds, in cypress ponds and bays and in grassy borders where
cypress bay met the open fields of Chesser Island.

Period. In 191 2 we found them breeding in May to July. In 19 14
(Deckert, 19 14, p. 5) at Jacksonville, Florida, "found their spawn, in shallow
bayous, in February, March, April, May, August and October." In 192 1,
basing our statements on 19 12 experience we wrote: "The southern leopard
frog, as well as the northern leopard frog, breeds normally in the spring, and
hence is called the 'spring frog,' but occasionally its breeding period may ex-
tend until July 4 or later, after the first eggs of the species are hatched and
the tadpoles transformed." In 1923 we (Wright, 1923, p. 34) held that
"From April 15 or earlier to September ist, six forms breed. . . . Rana
sphenocephala. . . ." In 1924 we (1924, p. 377) held "Species such as Rana
sphenocephala . . . which begin early in the season, breed during 25 to 30
weeks of the year, if not longer or from February to September or October."
In 1918 Pope (1919, p. 97) at Houston, Texas, on October 15, "Found three
clusters of hatching eggs, probably of this species, in a rain pool in woods
road." In 1920 Deckert (192 1, pp. 20, 21) in Dade County, Florida, writes
that "The writer has been expecting to hear the calls of the Southern Leopard
Frog, but none were heard during or after any of the heavy rains of May and
June. During the night of December 5, at Lemon City, after a violent thunder
storm the temperature dropped suddenly from 84°F. to 62°F., and when the
writer at 1 130 a. m. took a ramble around his yard with a lantern, he heard
the croak of what, according to his experience, must have been this frog.
The next morning was bright and quite cool, and the writer found, about
half a mile from his house, in a rain-ditch in a nearby rock-pit, three batches
of frog-spawn."

In 192 1 we found the southern meadow frog eggs from April 23 to July 23.
In 1922 we found them from June 22-August 9. These accumulated records
of others and ours give February to December for breeding. Doubtless
January also has its breeding frogs.

Temperature and humidity. Some of our records for ovulation are :

April 23, 192 1 May 21, 192 1 July 23, 192 1

April 24, 192 1 June 25, 192 1 June 22, 1922

May 13, 1921 June 30, 1921 July 13, 1922

May 16, 192 1 July 7, 192 1 August 9, 1922

Most of these were laid at night when temperatures range from 50-70°.
Our day records for these ovulation records of May- July, 192 1, and June-
August, 1922, are from 70-78° mainly for midforenoon or earlier. In late
April we recorded some air temperatures of 58-65° or lower. The discussion
for voice is pertinent to this topic. Humidity and excessive rains are very
important in determining onset of breeding.

Egg-laying process. In the field regularly and in camp usually they laid
plinth-like masses. We did not see the actual laying. Similarity of egg mass
form must make the process akin to that in R. pipiens. They do not move
from the stick until whole complement is laid.

428 Frogs of the Okefinokee Swamp

EGGS

Attachment. In 192 1 on April 24, at 9:30 a. m. we found "on the east
end of cypress pool in amongst pickerel weed attached to the stems is an area
of Rana sphenocephala eggs. Twelve masses, one fresh not yet expanded.
The eggs are from yg~2 inches below the surface of the water. Or, on June 6,
in another pond egg masses were "in the middle of a pond and around pickerel
weeds. Greatest number of egg masses yet. They are all submerged 2-6
inches below the surface. They are also around the edges of the pond as well
as being attached to other plant stems." On May 24 "Around Long Pond in
amongst lizard's tail (Saururus) are several Rana sphenocephala masses
hatching." In some of the smaller ponds they are attached to a variety of
plants and occasionally are on the bottom unattached.

In 1922 on June 22 at Camp Pinckney in water-filled pit Nos. 2 and 3
found fresh eggs attached to wild rice in the middle of the pits. In the road
just before Camp Pinckney found two or three masses, one unattached." Or
on July 13 we found them attached entirely to grass blades in a temporary pool.

Egg mass. In 19 12 in Okefinokee Swamp we found the egg masses of this
species in June and July. In 1918 Pope (1919, p. 97) at Houston, Texas, on
October 15 "Found three clusters of eggs, probably of this species, in a rain
pool in woods road. The clusters had evidently been round and about four
inches in diameter, though considerably softened and expanded by the hatch-
ing of the tadpoles."

In 1920, Deckert (192 1, p. 21) on December 6, 1920, at Lemon City, Fla.,
found "three batches of frog-spawn. The masses were partly afloat and at-
tached to weed stems in about 12 to 15 inches of water, milky with disin-
tegrated limestone. Securing one of the masses and placing it in a wooden tub
in our arbor, the eggs hatched on the 7th, and the tadpoles, though very small
as yet, can be positively said to belong to this frog."

In 192 1 on April 24, 192 1, we found as in 191 2 the "masses are plinthlike,
not globular. One mass photographed is i inch thick, 2 inches deep and 5 or
6 inches wide. Egg masses look much like those of Rana pipiens.^' In 1922,
on July 13 we noted "plenty of masses of eggs submerged. They are plinth
masses. The egg envelopes vary in size."

Egg description. On April 24, 192 1, we made these field measurements of
some eggs as follows: vitellus 1.5-1.8 mm., average 1.6 mm.; inner envelope
3.2 mm., outer envelope 5.4 mm. Later, Mrs. Wright in laboratory measured
these same eggs for illustrations and secured the following results : Of 2 7 eggs
the vitelli of two were 1.4 mm., thirteen 1.6 mm., and twelve 1.8 mm., the
range 1.4- 1.8 mm., mode 1.6 mm., average 1.7 mm.; of the inner envelopes
there were ten 2.4 mm., nine 2.6 mm., six 2.8 mm., one 3.0 mm., the range
24-30 mm., mode 2.4 mm., average 2.6 mm.; of the outer envelopes two were
3.4 mm., two 3.6 mm., ten 3.8 mm., thirteen 40 mm., range 3.4-4.0 mm.,
mode 40 mm., average 3.8 mm. A few inner envelopes were irregular or dis-
torted in shape and look as if due to shrinkage. In 1922 we made the field
notes of vitellus 1.5-1.8 mm., inner envelope 3.2 mm. In 1923 we gave the

Rana sphenocephala 429

egg complement of this species as 1054 from a field note entry of 1922. On
July 16, 1922, we made a count of two egg complements. "The total count of
the two complements combined is 2 117. These two were relatively small
complements. The complement of a large female might reach 1 200-1 500 eggs
or even more."

Dangers. Rapid drying of the ponds is a serious menace. On May 24,
192 1, we went to one pond which was drying up fast. The same night we
make this note: "Newt Pond much lower and two Rana sphenocephala masses
left high and dry. These laid a night or so ago. Even though the period of
hatching be short eggs attached to plant stems 2-6 inches beneath the surface
may be at the surface by hatching time from drying up of ponds. The name
Newt Pond above also suggests the one animal, the newt which relishes
frogs' eggs.

An amusing mistake is apropos. On May 27, 192 1, on Billy's Island in a
run we found "some yellowish eggs fastened to a stick. They bother me.
What are they?" Because we wanted to see and thought we saw strings (like
those in Amphiuma) between the eggs we called it Siren Run. Later on May
27, we queried "What are the yellowish white eggs? They are beneath the
surface. Flat plinthhke somewhat like R. sphenocephala eggs in egg mass
form. They are near the edge of the run. Another bunch near the north
edge and an area of 6 or 7 bunches on the south side. Shallow water. One
mass amongst moss near clean water's edge." Later the same day we found
"There were some 6 or 8 more masses of Siren other than those I first recorded.
In amongst iris several were .5-.33 of an inch below the surface. They are
very loose masses. They are yellowish white with no particular animal or
vegetable pole so far as color is concerned. They are in areas: one of 5 masses;
one of 4; one of 2; and thus it goes. Does each mass represent a female?
Did she wind a mass around the stem?" Still later, the same day we noted
"5 masses of Siren. Two or three inches below the surface. These are near
an extending bar of land with a pine tree as its center. About it are Leucothoe
bushes, etc. All of five are on the south side. Some masses attached to
Leucothoe branches lying in the water. Another area of four, two at bases
of growing iris and two fastened to old iris blades in water. Masses 1-3
inches below the water. Water coffee-colored, sphagnum comes into the water.
On the surface above these Sireji (?) eggs are eggs of Hyla femoralis. This
Siren (?) Run has small pines, Leucothoe, Iris, Sphagnum, Ilex glahr-a outside
and Ilex cassine sapHngs, sedges, I tea, Smilax in run. The water where
Siren (?) eggs are is at least 85°-9o°." Before June 3, we found several more.
Now we are satisfied they were not Siren eggs but spoiled R. spheno-
cephala eggs. There were as many as 40 or 50 masses in all, similar in color.
None of these in the field hatched to our knowledge. We concluded that we
were straining our eyes to find a string of jelly between the respective eggs
and then knew they were not Siren eggs. On June 3 the "run was getting
very dry. Some of the eggs are high and dry." Quite Hkely this happened
when the eggs were first laid and they were spoiled by drying. Subsequently
it rained to reswell the jelly. Still we are puzzled by one entry in our journal

43 o Frogs of the Okefinokee Swamp

that some of the eggs on May 28 hatched at noon in camp. If this be true
they are albinistic but why should 50 pairs of frogs lay albinistic eggs? Herein
doubtless lies another mistake on our part, a problem we did not entirely
solve.

HATCHING PERIOD

Eggs taken April 24, 192 1, 4:00 p.m. from a mass very fresh apparently
laid the evening of April 23-24 hatched "a day before April 28." On April
28, we make the entry: "Today some of the eggs of R. sphenocephala are
hatched. Taken April 24. Four or five days for hatching." The air tem-
peratures at night went down to 60° and by day up to 90° in this period. Of
the same eggs in the field we note on April 28 that "over in Ra7ia spheno-
cephala pond the eggs laid night of April 23-24 have been hatched a day.
Temperature of water at 4:00 p.m., 82°." In another pond one of the boys
found masses hatching April 28. They were close to the surface where the
water at 3:00 p.m. was 90°. On May 13 we found several masses and on
May 16 they were hatched and gone. Masses found May 21 were hatched
May 24. Other masses found on July i and June 30 were hatched and
hatching and not yet hatched on July 5.

In 1922 in the morning of July 13 at 12:30-1 :00 a.m. we heard a great chorus.
Visited the spot and found fresh masses. These eggs were ready to hatch
and hatching July 16. The air temperatures ranged from 68° at night to
88° at night in this interval.

We thus have 3-5 days for hatching in these April- July months. Possibly
in the late fall and winter months it may require almost as long a period as
for our northern R. pipiens.

MATURE TADPOLE

Color description from life {June 23, 1921). On sides of the body and over
the back yellowish olive or oil green, warbler green or olive green. The back
and sides are with conspicuous black spots. On the sides and over the
bronchial and pectoral regions is a mottling of light brownish vinaceous or
pale vinaceous drab, giving a bronzy appearance. The middle of the belly
is a solid color pale cinnamon pink, light vinaceous cinnamon or vinaceous
cinnamon.

Tail. The muscular part of the tail, upper and lower crests are with
larger black conspicuous spots than the body. The tail has some pale pur-
phsh gray, purple gray, pale violet-gray or violet-gray.

Iris black heavily punctate with greenish yellow dots; inner rim orange.

General appearance. Tadpole large (74.5 mm.) full and deep bodied.
Venter (Uke that of R. pipiens) not strongly pigmented so that viscera
clearly show through in life or in preserved specimens. Tail medium in
length, tip acute. Dorsal crest in width not equal to depth of musculature
and extending on to body somewhat ahead of the vertical of the hind legs.
Spiracle sinistral, decidedly upward and backward in direction, just below
axis of the body. Eye on lateral axis and nearer lateral outline in dorsal
aspect than mid dorsal line. Anus dextral, opening round or elHptical; mucif-

Plate XLI
Southern leopard frog (Rana sphenocephala)

1 . Siren Run, looking N. N. E. Billy Id., Ga. May 27, 1921.

2. Tadpoles, from white clay pool and normal habitat, Thompson's Landing, St.

Mary's River, Ga. July 13, 1922.

3. 4. Tadpoles, Chesser Id., Ga. July 25, 1922. x 1.2.

5. Tadpoles with four legs, Chesser Id., Ga. July 3, 1922. x 1.2.

6. Tadpole with stub of tail, Chesser Id., Ga. July 31, 1922. x 1.2.

7. Transformed frog, Chesser Id., Ga. August 11, 1922. x 1.2.

Rana sphenocephala 431

erous crypts more or less distinct in mature tadpoles and sometimes quite
distinct in half grown tadpoles. The dorsal row of either side starting near
the dorsal crest goes diagonally forward and outward to join the principal
lateral row which goes from above the middle of the insertion of the tail
musculature to back of the eye where a supraorbital and an infraorbital branch
goes forward.

Mouth parts: Teeth 2/3 or 3/3. Upper labium edged by a fringe of
teeth, the fringe longer than the beak. Below either end of this fringe is a
long row of teeth about 2/3-1/3 the length of the upper fringe. The median
space one half to one times the length of the lateral row. Usually it is less
than once the lateral row while in R. pipiens it is almost invariably more than
once the lateral row. In many tadpoles as they approach transformation
there appear in either corner a short third row of labial teeth on both sides
or sometimes on one side only. Also in mature tadpoles it not infrequently
appears. For example, in 19 12 we secured 23 tadpoles of this species, of
which six had the third row on either side, this row .33-.24 the length of the
second row and in three specimens the third row was present in only one side.
On the lower labium are three rows of teeth. The third row is shorter than
the single row of labial papillae, 1/3 to 2/7 shorter than the second or first
row, or about 1/6 to 1/5 shorter than the horny beak. First and second
lower labial rows about equal and slightly longer than beak. First one is
sometimes subdivided in the middle.

Measurements. Length of body (19.0-25.0) in tail (28.5-37.5 mm.)
1. 35-1. 66, average 1.53. Width (10.-16.5 mm.) of body in its own length
1. 56-2. 1, average 1.77. Depth (9.0-15.0 mm.) of body .8-1.36 in body
width, average i.i. Depth of body 1.5-2.3 in body length, average 1.95.
Depth (10. 5-16. 5 mm.) of tail in length of tail 2.0-2.66, average 2.65. Mus-
cular part (6.0-9.5 mm.) 1.35-2.35 in depth of tail, average 1.80. Spiracle
1. 2-1. 7 nearer base of hind legs or vent region (8.0-14.0 mm.) than the tip
of the snout (12. 5-16.0 mm.), average 1.46. Spiracle 1.1-1.86 nearer eye
(5.0-9.0 mm.) than base of hind legs or vent, average 1.46. Eye i.i 5-1.3
nearer to tip of snout (5.0-7.0 mm.) than to spiracle (5.0-9.0 mm.), average
1.2. Nostril to snout equal to nostril to both 3.6-5.0 mm. Mouth (3.0-5.0
mm.) usually 9-1.6 larger than internasal space (2.2-5.0 mm.), average 1.25.
Mouth contained 1.4-2.0 (average 1.68) in interorbital distance (5.5-7.5 mm.)
Internasal space contained in interorbital space 1.4-2.3, average 1.95.
The measurements of the largest tadpole are :

mm. mm.

Spiracle to vent 14.0

Spiracle to eye 7 . 5

Eye to snout 6 . 5

Eye to nostril 4 . 5

Nostril to snout 3 . 5

Mouth 4 . 5

Interorbital distance 7.0
Internasal distance 5 . o

Total length

62.5

Body length

25.0

Body depth

150

Body width

16.0

Tail length

37-5

Tail depth

130

Musculature of tail

9-5

Spiracle to snout

15-0

43 2 Frogs of the Okefinokee Swamp

General remarks. On April 28 we found mature tadpoles of R. spheno-
cephala. They are all in the same stage therefore not Rana clamitans. Other
reasons are : they have bronzy bellies like R. pipiens but the tail has prominent
black blotches unlike northern R. pipiens.

LARVAL PERIOD

Of this common frog we have little data on the actual larval period.
Eggs laid April 24, 192 1, and hatched April 28 were tadpoles with two large
hind legs June 17 and transformed July 4 or 71 days from egg deposition or
67 days from hatching. Others were transforming in some places July 23
or go days from laying. This gives 67-86 days for larval period. Much the
same period for the northern meadow frog. Frankly I believe the period
shorter in the south. Eighty-six days are probably too long for the period in
general and many may not live as much as 67 days as larvae. In 192 1 we
found this species transforming as early as April i to April 30. This means at
the "67-86 days" determination eggs laid as early as January 23 or January 4
or February 22 or February 3. In these winter months possibly earlier eggs
and longer period. In general I suspect 50-75 days may prove to be nearer
the true larval period.

TRANSFORMATION

Period. In 19 12 we found shortly after our entrance (May 25) into the
swamp, transformed specimens. The records are for May 28, 29, June i, 3,
6, 12, 24. In 1921 we recorded transformed and transforming specimens
April I, May 4, July 4. In 1 910 Professor J. C. Bradley secured them in
September and October. In 1922 we secured them transforming July 4 and
August 14. We thus have every month from April to October for trans-
formation. No doubt every month in the year some southern meadow frogs
transform though they are called "vSpring Frogs" as breeders.

Size. Under growth we concluded that 20-23 mrn- represented the range
of transformation. On June 24, 1912, on Billy's Island we found mature
tadpoles and transforming frogs. Twelve of the latter measured 18, 18, 19,
20, 20, 21, 21, 22, 22, 22, 22, 23 mm. respectively, the mode 22 mm., the
average 21 mm. On July 23, 192 1, on the same island we took 19 transformed
and transforming specimens in ditches beside the lumber railroad. They
measured 22, 23, 24, 25, 25, 25, 25, 25, 25, 25, 26, 26, 26, 26, 27, 27, 27-29, 30
mm. respectively, the mode 25 mm., the average 24 mm. The total range
of the two lots is 18-30 mm., the mode 25 mm., the average 23 mm. The
total range of our recorded specimens is 18-33 mn^-

General reinarks. In this species the black spots on the tadpole tail
before transformation are very conspicuous. In this regard they seem
different from our northern Rana pipiens tadpole (of Ithaca, N. Y.) just
before transformation.

This species shows considerable variation in transformation sizes. Why
one group should range from 18-23 mm., another from, 22-30 mm. and
another (not mentioned before) from 26-33 mm. is hard to explain. Are

Rana sphenocephala 433

local conditions of drying up, absence and plentitude of water or food respon-
sible, or are those eggs laid in January, February, or late fall less likely to
reach as large a transformation size as those laid in spring or vice versa?

In 1920 when we wrote that the southern leopard frog transforms during
June and July at .75 to i inch in length, the average being seven-eighths of
an inch we were speaking only of our May 2 8- July 4 experience in the Oke-
finokee Swamp in 191 2. We then suspected it might be any month of the
year but dared not reach beyond our known facts.

GROWTH

In 1920 we gave as the presumptive evidence for R. pipiens growth 18-31
mm. at transformation, 31-44 mm. for the first year, 44-52 mm. for the
second year and 52 mm. for the third year.

In 192 1 we secured of Rana sphenocephala on April 24, five examples 56,
57, 58, 60 and 73 mm., apparently two different growth years. From May
15-23, we secured examples as follows: 52, 58, 62, 64 mm.; 75, 81 mm. — two
groups. On July 5, we secured 67, 68, 70, 77 mm. specimens. Oh July 23
we took two examples of another growth group, 40, 44 mm. Of transformed
examples we had one, April i, 33 mm.; one May 4, 28 mm.; one, July 4, 26
mm. The 192 1 evidence reveals 26-33 mm. as transformation or just beyond
transformation; 40-44 mm. for ist year olds; possibly 52-64 mm. for 2nd
year olds, and 67-77 or 81 mm. for 3 year olds. In 1922 on July 4 and 5 we
took transformed frogs at 20, 20, 22 mm., and a female at 71 mm. In August
9-14, we secured one 22 mm. at transformation and a male at 56 mm. In
June (19th and 28th) we caught respectively a 53 mm. and 75 mm. specimen.
We, therefore, had a 20-22 mm. transformation group, 53-56 mm. group;
and a 71-75 group. In 19 12, from May 2 8th- July 15th, we secured trans-
formed frogs at 22, 30, 32 mm.; on May 28-29, two at 22 and 24 mm.; one
of 32 mm. June i; two 27 mm. and 30 mm. June 3; one 32 mm. June 6.
On June 12, we secured one of 27 mm., one of 37 mm., and one of 71 mm.
— three groups. On June 13-14 we secured one 50 mm. On June 24 we
secured four at 32, 33, 35, 37 mm., one of 49 mm., one 63 mm., — ^apparently
three growth groups. From July 15 to November 15 we received three 42
mm., 65 mm., 71 mm. in length. In 1913 a male captured December 26
measured 78 mm., and a female 76 mm.

On the basis of 1912, 1921 and 1922 collections it would seem that trans-
formation came at 20-23 mm.; that one year olds are from 35-50 mm.; that
two year olds are 52-65 mm.; that three year olds are 67-78 or 81 mm.

FOOD

No thorough studies have been made or published on this form. R.
pipiens has been studied by Surface, Drake, Frost, Munz and Klugh. No
doubt the two species are quite similar though the range of the southern
meadow frog might cause its diet to be quite different.

After the above was written we chanced on Force's (1925, p. 26) para-
graph which follows :

434 Frogs of the Okefinokee Swamp

"Of the 46 specimens of Southern Leopard Frog, 10 had empty stomachs,
while the remaining 36 had 38% Elateridae, 16% IMyrmicidae, 11% larva,
4.5% each Lachnosterna and Coccinellidae, 1.8% each Saldidae, Belos-
tomidae, Scarabaeidae (Tumble Bugs and June Bugs), and Cucujidae, 2.7<>^
each Formicidae and Dytiscidae and .g% each Lamiinae, Cicindellidae,
Membracidae and Crayfish."

ENEMIES

On May 6, 1921, I found a cypress pond dried up except for a small
center of water filled with fish and R. sphenocephala tadpoles. Here Great
Blue Heron and Florida Crackles were having a fine feast. On July 7, 192 1,
we heard subdued croaks of a frog and soon found that the southern water
snake (N. sipedon Jasciata) had a meadow frog which it swallowed in 7-10
minutes. We photographed a water snake with a meadow frog in its mouth.

In 19 1 2 we found these results. A black snake (Coluber constrictor con-
stricto?') had swallowed one adult meadow frog. A pied water snake ( Matrix
taxispilota) had two meadow frogs in its stomach while two southern water
snakes (iV. s. jasciata) had each a meadow frog. One ribbon snake {Tham-
nophis sackenii) had swallowed two meadow frogs, and one snake of this
species had one meadow frog in its stomach. The true water moccasin also
preys on meadow frogs.

AUTUMNAL DISAPPEARANCE

(See First Appearance). Our records of autumnal disappearance mean
little for none of us have been in the swamp at the proper autumnal period
to record it. Strictly speaking there probably is activity the year through
as recorded under First Appearance.

AFFINITIES
Cope first recognized the southern meadow frog as a subspecies of R.
virescens. Miss Dickerson 1906 (p. 188) by fine illustrations and descriptions
did more to bring out the characters than any other author. Of it she writes:
"The species is evidently a very distinct one, not intergrading with Rana
pipiens, but holding its own with the latter frog in the same localities in the
southern part of the United States." It is not questionable whether there is
such a mixing of ranges. This form seems a Lower Austral offshoot of Rana
pipiens extending from New Jersey to Texas and up the Mississippi to In-
diana. We have treated this form as if it were very distinct from Rana pipiens
but the whole Rana pipiens group is a complex one.

BIBLIOGRAPHY

1920 Barbour, T. Copeia, July 31, No. 84, p. 56.

1925 Brimley, C. S. and W. B. Mabie. Copeia, February 15, 1925, No. 139, p. 15.

1920 Boulenger, G. A. Proc. Amer. Acad. Arts Sciences, August, 1920, Vol. 55, No. 9,

PP- 434. 436, 439-

t, R. Copeia, l*ebruary 14, 1914
1921 . Copeia, March 15, 1921, No. 92, pp. 20, 21.

1914 Deckert, R. Copeia, February 14, 1914, No. 3 and April 15, 1914, No. 5.

1922 . Copeia, November 20, 1922, No. 112, p. 88.

1906 Dickerson, M. C. The Frog Book, pp. 186-188.
1925 Force, Edith R. Copeia, April 30, 1925, p. 141.

Rana lurgatipes 435

1922 Loding, H. P. Geological Survey of Alabama. Mus. Paper No. 5, pp. 20, 21.
1924 Myers, G. S. Copeia, June 30, 1924, No. 131, p. 60.

1921 Ortenberger, A. I. Copeia, October 15, 1921, No. 99, p. 75.
1927 . Copeia, April-June, 1927, No. 163, pp. 46, 47.

1919 Pope, Philip H. Copeia, December 31, 1919, No. 76, p. 97.

191 8 Viosca, Percy. Jr. 3rd Biennial Rept. Department of Conservation Louisiana
April I, 1916-April I, 19 1 8, p. 160.

1920 Wright, A. H. U. S. Bur. Fisheries Doc. 888, p. 23.

1923 • Copeia, February i, 1923, No. 115, p. 34.

1924 Wright, A. H. and A. A. Wright. Am. Naturalist July-August, 1924, Vol. LVllI,
P- 377-

Rana virgatipes Cope

(PI. Ill, Fig. 3; IX, Fig. 2; XI, Fig. 13; XIV, Fig. 4; XVI, Fig. i; XVII; XLII; XLIll;

Text Fig. I, 14)

COMMON NAMES
Sphagnum Frog. Carpenter Frog. Cope's Frog.

RANGE

Check list. "Range: New Jersey to North Carohna. Type Locahty:
Near Atlantic City, New Jersey." Stejneger & Barbour (191 7, p. 39).

Supplementary records. The North Oarohna record no doubt referred to
Lake EUis records by C. S. Brimley (1906) 1907 ((1907, p. 159; 1909, p. 133)
and by J. Hurter (1907). Dunn (1918) reported it from Norfolk, Va. Wm.
Palmer in 1896 secured it at Lake Drummond, Va. Myers (Copeia, 1924,
p. 60) reported two from Wilmington, N. C. In 1922 (1923) we reported
the extension of its range from N. C. to Okefinokee Swamp, Georgia. In

1922 our party found it and Hijla andersonii at Everetts Pond which is .75
or 1.5 miles from the N. C.-S. C. state line. In June 1920 W. T. Davis and
Dr. J. Chapin (1922, p. 74) recorded it at Southern Pines, N. C. about
35-50 miles northeast of Everetts Pond. Noble in 1923 (p. 422) remarks
"Why R. virgatipes has a more extensive range to the south than H. ander-
sonii it is impossible to say at this time." Previous to 192 1 one would have
held H. andersonii farther south at Anderson, S. C, and farther north
in N. J. (per Miller) than R. virgatipes. We found both near the N. C.-S. C.
state line (near Cheraw). I believe them more or less associates and confi-
dently expect H. andersonii to be found in Georgia.

Local Okefinokee records. In 192 1 we recorded it first near Bear House
and Hurst Island (near Chesser Island) on July 12 and 13. In the ist and
2nd dreens or causeways between Chesser's Island and the mainland we heard
it from July 15-19 in the wooded swampy parts. Ten days later in a grassy
bay or head near Black Jack Island we heard it again. In 1922 we found it
around the borders of Chesser Island from June 21 until August 11.

GENERAL APPEARANCE
A small frog with long narrow head, back brownish with four yellowish
or golden-brown, longitudinal stripes on the back. The under parts are yel-
lowish white with brown spots. The femur has alternating dark and light
stripes.

436 Frogs of the Okefinokee Swamj)

MEASUREMENTS

(Recent Material)

Head to angle of mouth i.o (28 mm. near transformation)-:. 04 (36 mm.
9)-i.o8 (36 mm. cf)-i.o6 (44 mm. cr')-i.2 (56 mm. d^)-i.o (56 mm.
9) in width of head; head to rear of tympanum .87-.92-.96- 95-857-95
in width of head; head to angle of mouth 2.8-3.13-2.9-2.5-3.75-3.0 in length
of body; head to rear of tympanum 2.43-2.77-2.57-2.5-2.66-2.8 in length
of body; snout .Q5-1.18-.92-.94-.875-1.23 in fourth finger; .76-1.18-1,15-
.94-875-1.3 in first toe; eye i. 3-1. 57-1. 3-1. 23-1. 23-1. 21 in snout; .61-85-
.7-.69-1.07-.71 in tympanum; 1.12-1.85-1.2-1.0-1.07-1.5 in first finger;
tympanum 3. 2-3. 5-2. 7-2. 6-1. 8-1. 64 in intertympanic width; 2.1-1.83-1.71-
1.77-1.14-1.7 in snout; internasal width .66-85-1. 0-.66-1.0-.9 in upper
eyelid width; interorbital width 1.0-1.5-1.15-1.0-2.25-1.8 in upper eyelid
width; 1. 5-1. 75-1. 15-1. 5-2. 25-2.0 in internasal space; 4.0-5.25-3.15-4,0-
6.25-5.4 in intertympanic width.

Forelimh: 1.7 5-2.0-1. 9-1. 9-2. 43-1. 93 in length of body; 2.75-2.6-2.65-
2.67-3.2-2.62 in hind limb; first finger i. 33-1. 3-1. 33-1. 46-1. 3-1. 23 in third
finger; second finger 1.5-1.3-1. 33-1. 35-1. 3-1. 44 in third finger; 1.12-1.0-
1.0-1.0-.93-1.72 in first finger; third finger i. 08-1. 17-1. 37-1. 26-1. 44-1. 2 in
second toe; fourth finger .80-1.0-1.25-1.0-1.16-1.04 in first toe; i. 2-1. 37-
1. 33-1. 26-1. 5-1. 24 in third finger; internasal space i. 5-1. 85-1. 7-1. 44-1. 55-
2.1 in first finger; 1.33-1.85-1.71-1.55-1,55-1.8 in second finger; 2.0-1.8-
2.28-2.11-2.0-2.6 in third finger; 1.66-1.85-1.7-1,66-1,33-2.1 in fourth
finger.

Hindlimh: length 1.5 7-1. 3-1. 4-1. 3 2-1. 34-1. 3 5 in hind limb; tibia 2.07-
2.4-( )-( )-2. 43-2.33 in length of body; 3-25-3-i3-( )-( )-3-26-3.i6 in hind
limb; i.i8-i.2-( )-( )-i. 0-1.20 in forelimb; i.ii-i.33-( )-( )-i.o8-i.i5 in
hind foot; first toe 1.61-1.53-1,46-1.6-1.85-1.4 in second toe; 2.25-2.0-1.93-
2.26-2.55-2.0 in third toe; 3.0-2.6-2.6-2.93-3.43-2.45 in fourth toe; 2.0-
1. 84-1. 73-2. 0-2. 28-1. 82 in fifth toe; second toe i. 38-1. 3-1. 32-1. 41-1. 38-1. 4
in third toe; i. 84-1. 7-1. 72-1. 83-1. 83-1. 74 in fourth toe; 1.23-1.2-1.18-1.25-
1. 23-1. 29 in fifth toe; third toe i. 33-1. 3-1. 3 i-i. 3-1. 33-1. 25 in fourth toe;
.88-.92-.90-.88-.88-.93 in fifth toe; fourth toe i. 08-1. 35-1. 31-1. 54-1. 04-1. 5
in hind foot; i.i2-.88-( )-( )-.95-.9o in tibia; i. 25-1. 06-1. 0-1.04-.95-1. 07 in
forehmb; fifth toe i. 5-1. 4 i-i. 46-1. 46-1. 5-1. 3 5 in fourth toe; internasal
space 1. 33-1. 85-2.08-1. 66-1. 55-2. 1 in first toe; 2. 18-2. 85-3. 14-2. 8-2. 88-3.1
in second toe; 3.0-3.7-4.14-3.8-4.0-4.3 in third toe; 4.0-4.85-5.4-5.0-5.33-
5.4 in fourth toe; 2.66-3.4-3.7-3.3-3.55-4.0 in fifth toe.

HABITAT

Davis found (1905, pp, 795, 796) that "Rana virgatipes Cope is also more
abundant at Lakehurst than at first supposed and has been found from May
to September. In the early summer as many as twelve have been seen in
one day without much search having been made for them. They are apt to
be in some ditch and seated on the sphagnum moss, or on a floating lily pad,

Rana virgatipes 437

and they are found also in the ditches that border and intersect the cranberry
bogs, as well as in the lake."

The other extended account of their habitat is given by Fowler (1905,
p. 662, 1906 (1907) p. 123). "Our specimens were obtained at the mouth of
the Mare Run, the type locality, and we noticed no other amphibians in this
vicinity. Associated in this locality were many examples of Chnjsemys pida
and Chelopus guttatus, which were constantly bobbing around in the sphag-
num. Enneacanthus obesus, Mesogonistius chaetodon, Erimyzon sucetta ob-
longus and Notropis chalbaeus abbottii were also abundant. The frogs seem
to occur exclusively in the almost submerged masses of sphagnum which Hne
the shores in many places and often extend well out in the stream. Here
the water is still and the animals rest more or less below, so that their dull
colors harmonize well with the surroundings. They were shy and sank
quickly out of sight among the aquatic vegetation on the approach of danger.
In attempting to escape they would not jump or leap and when caught in
the dip-net moved about in a rather slow stupefied manner. They swam for
short distances, but were usually able to find suitable shelter close at hand."

Barbour (191 6 p. 5) "in search of the Sphagnum Frog, Rana virgatipes
Cope" at Lakehurst, N. J., writes:

"The afternoons we have usually spent in wandering about the sphagnum
bogs and along the cold spring branches which run into the lake, searching
for the sphagnum frogs, which we have found at times in considerable num-
bers. Our observations have confirmed those of Davis and of other observers
who have recorded their shy ways and strictly aquatic habits."

For our habitat notes of 192 1 see "Voice." In 1922 we camped the night
of June 9 on the north side of Everett's Pond (about 5 or 10 minutes by auto
from N. C.-S. C. state line), N. C. "In the wooded sections or streams
flowing into the lake in dense thickets of Smilax (briar) and other plants
were Rana virgatipes croaking quite generally. Several in different places."
The Hyla andersonii were in a wooded stream or branch swamp which flowed
into the wooded R. virgatipes area near the lake.

In 1922 on June 21 in the prairie where we first found the species in
Okefinokee and just north of Lake Sego (Seagrove) "heard 7 or S R. virgatipes.
Found one in a grassy place with waterlilies. On a waterlily pad was the one
I captured. They like grassy places about or near the edge of a head (islet).
Also seem to like some timbered parts or be in timbered parts entirely."

On May 22-23, 1924, we heard a few males croaking on the south side of
the west point into the lake at Lakehurst, N. J. "It was at noon with the
sun shining. Here in the sphagnum-heath edge caught 4 adults, two females
and two males. As we waded along we would see them sometimes wholly out
of water or at the lake's edge. Usually they leaped into the water and hid
under the vegetation mat or quickly came up under a water fily pad or swam
some distance and then poked out their heads. Most of the males were farther
out in deeper water. The heaths and birds such as prairie warbler, pine
warbler, white-eyed vireo, towhee, bob Vv^hite, turkey buzzard, etc., suggest a
southern environment." On May 23 from the west point northward "the

438 Frogs of the Okefinokee Swamp

west edge of the lake where the Rana virgatipes are normally common had
the sphagnum just starting and below the top of the water. Pipeworts
(Eriocaulon) were just starting. Water lily leaves were yet below the water.
Golden club (Orontiuni aquaticum) was in bloom but we saw no adult frogs in
this stretch. Some little frogs just ahead of me were continually disappearing.
They were probably transformed R. virgatipes but of this I could not be sure."

In 1929 in June we planned a trip to Lakehurst, but a mishap befell us.
Of the improvised trip June 8, 1929, we have these journal notes.

"Needed a male and a female R. virgatipes for photographs. Boarded a
train at Ridgewood for Lakehurst and without collecting kit or clothes. It
rained until midaf ternoon ; with every handicap we waded the edge of the
lake at Lakehurst with no success. The same result along the west point of
land extending into the lake. The vegetation mats have not reached the sur-
face to any appreciable degree. Mr. Shinn and his boys have a cottage at the
base of the point. They informed us that locally the frogs have long been
called 'carpenter frogs.' Some new visitors beside the lake can hardly sleep
at night when the species is in full chorus. Mr. Shinn sent the boys out in a
canoe to get some on the mats in midlake and along the east side. We went to
Emlie's pond. When we returned the boys had 4 males and i female and
one recently transformed frog. Several they had thrown back. They ap-
proached the Hly pads and with a scoop of a long handled dip net caught the
frogs. The lily pad and mid-lake habits suggest Ra7ia septentrionalis of the
Adirondack Mts. and the North."

Outside or inside the swamp. Rana virgatipes in 192 1 we missed in the
center of the swamp though one informant claims he has heard them on the
south side of Billy's Island. They are not in the center of the swamp in our
experience. We found them on the prairies near Chesser Island and between
it and the mainland. Also we recorded them from Blackjack Island. This
places them in the very southern and eastern part of the swamp, a queer dis-
tribution as puzzling as that of the plant Bejaria (Elliottia) racemosa.

FIRST APPEARANCE

In the Okefinokee swamp some of the residents to whose attention we
called this frog, said it started croaking in early spring if rainy weather be at
hand. They probably appear by mid- April if not before.

VOICE

According to Davis (1907, p. 50) "Rana virgatipes may be called the Car-
penter Frog, for its note sounds much like the blow of a hammer on a board.
It is a quickly uttered chuck-up, chuck-up, and the frog usually hammers three
or four times. For a time I was not sure of the singer, but some captive indi-
viduals under the influence of good living have uttered this call-note in my
room while I sat by,"

In 1907 Fowler writes (1905, pp. 662, 663; 1906, (1907) pp. 123, 124, 125):
"The individuals obtained by Cope and Stone did not make any noises, but
when we discovered ours, the males were in full cry, which would seem to indi-

Plate XLII
Carpenter frog (Rana virgatipes)
Second "Dreen," near Chesser Id., Ga. July 22, 1922.

2. Egg mass, Chesser Id., Ga. July 22, 1922, x i.o.

3. Eggs, stained, Chesser Id., Ga. July 22, 1922. x 1.2.

4. 5. 6. Male croaking, second "Dreen" near Chesser Id., Ga. July 21, 1922.

Flashlights.
Male, Chesser Id., Ga. July 15. 1921. Lateral aspect, x 0.4.
Female, Chesser Id., Ga. July 22. 1922, Lateral aspect, x 1.2.

Rana virgatipes 439

cate that late April was the height of the nuptial season. The males are pro-
vided with distensible vocal vesicles, and when about to utter their call these
sacs are inflated like httle bladders till nearly spherical, and then by degrees
the air is allowed to escape. This gives the sac the appearance of collapsing
by a series of jerks. As it is done quickly, each jerk at an interval of a second,
the result is a sort of a rapping sound. These raps or jerks are about five or
six in number. The sound produced is peculiar in that it is difficult at times
to detect its source, and if the frog is close is quite startling. This is due
not only to the suddenness, but also to the volume of the sound. On one occa-
sion an example which I captured has his vocal vesicles well inflated, and
though they partly collapsed as I held him in my hand he did not utter any
sound. The call bears considerable resemblance to the sound produced by
wood choppers cutting trees a short distance back in the forest, and is different
from the cry of any other batrachian, so that when first heard I suspected it
was produced by this species."

"Mr. J. A. G. Rehn informs me he heard this frog at Cedar Grove, on the
Oswego River, or the east branch of the Wading River, and also at Speedwell
during June of 1905. A description of the cluck of a frog, which may possibly
have been this species, was given to me by Mr. G. L. Hartman of Palermo,
Cape May County, who says that the frogs are common about the Cedar
Swamp Creek region. Although diligently searched for in all suitable locali-
ties I failed to discover any. Mr. Chreswell J. Hunt informs me that he noted
this frog while camping on the dam of the Rancocas Creek at New Lisbon, in
Burhngton County, on the night of May 15, 1906. During all the evening its
voice was the sound most in evidence. It was also abundant at the dam at
Brown Mills on May i6th. About the shores at the head of these ponds were
masses of sphagnum and in these spots the frogs seemed to abound. Although
unsuccessful in securing specimens, he feels certain of the identity of the species
as the croak was so different from that of any other with which he was familiar.
He also states that he did not hear it anywhere along the Rancocas below
New Lisbon, which is well among the pines. During the past summer he
heard it at Whiting. On May 30, 1907, Mr. Percy Lorrilhere reports he heard
this frog at Dennisville in Cape May County." Again in 1907 Fowler (1908,
p. 194) gives us the following: "Rana virgatipes Cope — Sphagnum Frog. The
clack, clack, clack, clack, clack of this frog could be heard about the Rancocas
Creek above New Lisbon, in Burhngton County, on May 12th, 1907. It
occurred at intervals, interrupting the stillness of a backward spring. Though
loud and not often uttered, in comparison with those found at Mare Run, their
croaking could be heard at quite a distance. Sometimes the animals must
have been quite close, for we could hear them when but a few feet away with-
out seeing them. It may have been that the weather was too cold, for their
croaking was always located as coming from among the submerged and over-
grown vegetation along the banks. In such places the temperature was con-
siderably higher by mid-day than elsewhere. When Mr. Hunt visited this
place just a year previously, the frogs were very numerous. The weather at
that time, however, though about the same time in May, was much warmer.

440 Frogs of the Okefinokee Swamp

In the evening, during the night, and in the early morning, they were very
noisy, but during the day were more or less quiet, only an occasional croak
being heard at intervals, or as noted above."

Our first record for Rana virgatipes came July 12, 192 1, near Chesser
Island. We had had to abandon our boat because of low water and wade to
shore. Near a head not far from the boat about 5 p. m. we began "to hear a
new Rana calling in the maiden cane — cluck-cluck-cluck-cluck; a rather loud,
incisive, distinctive, clucking croak." (F. Harper). The air temperatures
were at least 75°. On the following morning (air temperature about 78°;
cloudy day) the author with four of the Chesser boys "went for the rest of
the equipment at Bear House (af miles from Chesser Island). Heard no end
of Rana clamitans, R. grylio and one or two of the unknown {R. virgatipes'!).
Also over toward Hurst Island heard several (R. virgatipes'!)". The previous
afternoon we instinctively pronounced it R. virgatipes though we had never
heard it but had read previous characterizations of it. On the night of July
1 5 after a thunder shower we heard several in the wooded dreens or causeways
between Chesser Island and the mainland. Temperature about 70-73 degrees.
On July 19 we heard a few before dawn towards the open prairie and at night
we caught another in the wooded causeway. On Black Jack Island July 28,
1921, we had one of the hardest rains I ever experienced. In the morning of
July 29, 192 1, when the temperature was 73 degrees "I heard a Carpenter
Frog chorus. . . . Just outside the south bushy edge of Black Jack Island
(near camp) were the carpenter frogs. They were scattered in among the
small cypress where there was an undercarpet of Woodwardia and a little
sphagnum. They seemed to be around the base of the cypress trees and on
the cypress knees. I saw none of them. In the daytime they are shy. The
croak is 4-7 notes in rapid succession, all alike. Then the species has another
call of two notes and lower in pitch." Later "on the sphagnum strand many
roads out from Chesser Island we heard some carpenter frogs." The period
from July 12-19 was a very rainy week throughout southern Georgia and no
minima of nearby stations went below 69°, our lowest temperature being 73°.
On July 28 almost 3.80 inches of rain fell at Waycross and about 2.4 inches the
next day and the temperature ranges from 7o°-84°.

In the Okefinokee swamp we heard them at different times of the day.
Summarized our 192 1 record reads somewhat as follows:

June 19. Heard at night in wooded sloughs.
" 24. ,"I awoke at 3 a. m. and heard quite a few calUng towards the edge of

the prairie."
July 2. Harper heard one on prairie at 8-8:30 a. m.
" 21. Calling generally in wooded sloughs.

" 22. One heard in mid-afternoon (2:30 p. m.) in sloughs. Sunlight.
" 22. At night many croaking in slough at temperature 72° until midnight

or later. Other frogs not active.
" 25. At night many croaking in slough. Temperature 7 2°-74°. Air humid.

Croaking beyond midnight. Other frogs not active.
" 28. At 3-4 a. m. heard several. At 6:30 a. m. heard one. Air 72° cloudy.

Ra7ia virgatipes 441

Aug. 8. Heard two at 8 a. m. (Temp. 79°).
" 9. At 3 and 4-5 a. m. heard several on prairie or at landing. Heard at

night in bay.
" 10. Heard at 7.
" II. Calling at midnight.

In other words, they start about dusk or sometimes before and gather
strength of chorus as midnight approaches. At 3 or 4 in the morning they go
strong. At daybreak one to several may be heard and these may continue
until 8:30-9 a. m.

In 1922 on June 21 (Okefinokee Swamp) we have the record of another
call. "Male has two prominent lateral pouches. In this respect differs from
Rana grylio. When held in hand it will open its mouth and make a peculiar call.
Often it will swell up when held just as many a frog will when seized by a snake."

On the night of July 21, 1922, when the frogs were actually breeding we
made these observations: "Whenever one approached within 5 feet or more
usually the male stopped croaking. Seldom indeed did the male croak after
the light was put on it. Anna found an area where she heard a scrambling.
One croaked under observation. We set up the camera. It moved on. In
this area about midnight we secured three exposures. They are strangely
silent and wary when one approaches. The croak may be of two parts, some-
times three, four or five parts. Each part occupies a second or less in duration.
This species has a pouch on either side. These are round vesicles perfectly
round when the frog is croaking and the throat does not inflate. Therein as
before remarked they differ from Rana grylio^ Rana clajnitans and Rana
catesbemna. In this regard they are more more like Rana sphenocephala?
Are its eggs to be in a film or a mass like Rana sphenocephala? Doubtless the
former." We made the wrong guess. The next morning revealed the answer.

Our records for 1921-1922 indicate a range of croaking from June 9-Aug.
II. Mr. Benjamin Chesser of Chesser Island says "they croak in the spring
and summer alike during wet weather" and his father, R. A. Chesser, con-
sidered it one of the first frogs to begin croaking in the spring.

On May 22, 1924, from Mr. Emlie's pond, Lakehurst, N. J. we "heard an
immense chorus at Lakehurst lake. It began just before dark." "R. virgatipes
is more of a night frog. Once in a while we heard it at day in cloudy weather."

F. Harper makes these notes "call 2-5 times in series. Occasionally a
minor single note (between or preparatory to series.) Good loud clear note.
Frog sprawled in water like R. sphenocephala, or sitting half submerged, like
R. clamitans or perhaps only | under. Seldom less than about 15-30 seconds
between calls, intervals apparently irregular. Generally a silent premonitory
gulp. No swelling of throat but of lateral sacs, each to about h head. Between
each note collapse to about ^ size (small marble). At full inflation about
size of small persimmon or larger marble. Between series collapse entirely."

MATING

Male (From life, July 15 1921). Stripe down either side from eye to hind
leg hazel or fawn color. Back natal brown or chestnut brown. This color
with black spots on the back. Natal brown on dorsum of fore and hind legs.

442 Frogs of the Okefinokee Swamp

Same color with black on sides. Smaller stripe from angle of mouth halfway
on side Hght or pale ochraceous salmon. This stripe continued over angle of
mouth and on to upper jaw pale greenish yellow. Throat sulphur yellow with
indefinite brown spots. Rest of under parts cartridge buff to pale chalcedony
yellow with beautifully marked bone brown or almost black spots on venter
and sides. Back and fore parts of iris neva green. Iris inner rim green and
yellow. Iris proper spotted black and empire yellow.

Female {From life, July 22 1922). Back stripe buffy brown and side stripe
also. Back color olive brown. Throat sulphur yellow, also inner edge of sur-
face of fore limb. This yellow also from angle of mouth to over arm insertion.
Spotting on under parts not so prominent as in male nor so clear cut yet it is
conspicuous. Under parts and sides with hardly any wash of yellowish as the
male sometimes has on his sides. Females in general on under part look white
while males have washes of yellow on throat, sides and slightly on other parts.
Front edge and back edge of eye bright green yellow as in the male (see male
of July 15, 192 1 ). This female smaller than the four males captured at the
same time.

Female {From life, July 25, 1922). This gravid female has as dark a back
ground color as a male and in this respect unlike above specimen of July
22, 1922.

Structural differences. In 1906 (1907, p. 121) Fowler notes the males have
"Vocal vesicles large and capable of considerable inflation" and presents a
striking plate of the distended sacs of the males and the venters of two males
and a female. Boulenger (1920, p. 430) says the male has "a large grayish or
blackish external vocal sac on each side between the mouth and shoulder; a
feeble pad on the inner side of the first finger."

The males have the tympanum enlarged but not so strikingly as in Rana
grylio, Rana catesbeiana, Rana clamiians or Rana onca nor is the thumb quite
as large proportionally. I cannot, however, call it a "feeble" pad.

The young males at 25-39 mm. have thumb Httle swollen, tympana not
very large and vocal sacs not well developed. About 40 mm. the thumb be-
gins to enlarge, vocal sacs begin developing and tympana may or may not
be large. Certainly by 45 mm. if not before, thumb, vocal sacs and tympana
are well developed in the males. However one male at even 38 mm. begins to
have thumb enlarging and vocal sacs developing and one male at 42 mm. is
almost a full male in tympana, sacs and thumbs.

Duration. Probably of short duration normally.

Night or day. Mainly by night, though by day in cloudy or rainy weather
when the climaxes come in breeding.

Amplexation {normal, abnormal, cross). It must be pectoral in this species.
No mated pairs have been secured. This species is shy like Rana grylio and
Rana catesbeiana. Unlike toads and tree toads none of the males would seize
the gravid females we placed with them.

Rana virgatipes 443

OVULATION

Habitat. In 1923 (p. 421) Noble lists the breeding habitat of Rana
virgatipes as "Larger pools of pine barrens." In 1908 Fowler (1908, p. 194) in
writing of this species "above Rancocas Creek above New Lisbon, in BurHng-
ton County" says "We found a lot of spawn, though were unable to identify
it as belonging to this species. It resembled that found at Mare Run, in
similar situations, as it was formed in strings and wound about the numerous
aquatic plants in still water. These strings were quite long and very abundant.
Attempts at their transportation and hatching failed." These strings do not
accord in description with our experiences with R. virgatipes eggs. In 1922 on
July 2 1 we heard several Rana virgatipes in the wooded swampy areas between
Chesser Island and the mainland. "At 9 p. m. we started with auto for the first
and second dreens. At the first heard a few Rana virgatipes males calling. In the
area around the pond south of the first crossing were most of the frogs. Other
species not caUing in great numbers, so we went to the 2nd crossing. To
the direct right in the wooded edge of the island we heard several males.
These males would be next to a sphagnaceous base of a pine or cypress or
between two bushy clumps or sedgy tussocks. Occasionally they were perched
on a log. Sometimes they were amongst Xyris or Eriocaulon stands. There
were fallen logs in this area. The water at the deepest was no more than a
foot. The places of gathering were near the edge of the wooded swamp.
This species seems to be very appropriately a 'sphagnum frog.' There must
have been twenty or twenty-five frogs in one small area. We captured four
males and saw 10 or 12 more."

The next morning at 11 a. m. M. D. Pirnie and I went to the 2nd dreen to
look for the eggs. We found six masses of eggs. Our rough notes describe the
area thus: "The area has pine, cypress (young) and gum. More or less open
spaces between trees. Around the bases of the trees are sphagneous mounds.
More or less wide clear interspaces of water fill most of the rest of the area
except for smaller stands of Eriocaulon, Xyris, Poly gala cymosa, and Smilax
Walteri. The Eriocaulon stands may have Burmannia, Drosera with long
scapes and white flowers. Some algae and two bladderworts, one purple, one
yellow in water interspaces. Sometimes a stand around a cypress or pine is
Rhexia mariana or Woodwardia. Little of latter. Interspersed with trees are
Hyperica-like bushes of very fine leaves ^-f of an inch in length. Clethra
here and also evergreen "hurrah bushes (Leucothoe) ." Later in the day we
found more egg masses.

Period. In 1923 Noble (p. 425) considers Rana virgatipes a protracted
b»eeder, temperature controlled, a form which has hibernated in the water.
In Okefinokee Swamp and throughout most of its range I fancy rains and
humidity are more important factors than temperature though the latter is
an influence as well. We actually found ovulation from July 21-25 but we
have presumptive evidence of breeding from June 21-Aug. 11. Probably the
species begins the last of April and extends to August.

Egg-laying process. We have not observed the process. It is doubtless
similar to that for forms which lay beneath the surface such as Rana pipiens,

444 Frogs of the Okefinokee Sivmnp

R. sphenocephala, R. aesopus, R. septeritrionalis, R. syluatica, and other be-
neath-the-surface layers. The process probably requires a few minutes. The
female may start slowly but once the process begins it doubtless is over in
10-30 minutes.

EGGS

Attachment. The masses found July 28 were attached as follows: The
first mass was "found in a clear area attached to a Xyris blade and pine needles
and beneath the water, ..." Six more were found the morning of July 22,
1922. "They were attached to pine needles, upright sticks, Xyris or Erio-
caulon stems." In the afternoon we "found a mass much farther west than
the original area of 5 masses. It looked blue above and when turned over
showed a creamy white impression. The mass like those of Rana spheno-
cephala and Rana pipiens, looked white below. The mass was 2f x if inches.
As in Rana sphenocephala and Rana pipiens it seems to lay in groups as in
case of 5 masses yet this last fresh mass was considerably apart from others
in water 6-8 inches deep. If most of the eggs are laid beneath the algae on
the surface I am lucky to find these few masses. The species may, however,
choose clear water areas with little algae for ovulation." Other observations on
attachment are "Attached to fresh floating pine needles (Pinus carihaea). . . .
Attached to rosettes of Eriocaulon |-f inches below the surface."

Egg mass. The masses were more like the eggs of Rana sphenocephala
than those of Rana grylio. The first mass was "not globular but somewhat
elongate. The top of the mass was at or just beneath the surface of the water.
No rain had intervened since last night. The mass may have been 1.5 x 3 to 4
inches." "About eight or ten feet farther into the area were four more masses
two not far apart and two very close together. Isolated was another mass.
These five with original mass found makes six. These were all beneath the
surface. Very small masses for frogs. The temperature when we found them
was 72°."

Other data is as follows: "Sometimes in looking for Cope's frogs' eggs we
mistake the bladders of Utricularia purpurea for a mass." "... A mass
2x2x2 inches." "Another mass in water 4 inches deep, i inch below surface,
I inch thick, plinthlike, 2I x 2I inches, a perfect duplication of R. spheno-
cephala masses. This last mass 15 feet from two others which are 4 feet apart."
"Another mass, water 3 inches deep. Beneath surface attached to Eriocaulon.
Very much like southern meadow frog's mass." "One mass attached to Erio-
caulon leaves, if x 4 x i inches, a fine plinth but elongate, strap-shaped."
"Another mass attached to Rhexia mariana."

One has to be careful. "Right near one of the Rana virgatipes mass was a
very large mass as big as some Rana sphenocephala masses. Is it R. spheno-
cephala? We took it in to camp. Later in the day we proved it to be Rana
sphenocephala. The eggs were different and had inner envelopes."

Egg description. Our first field description of the egg is "The eggs are
large, far apart like Rana sylvatica eggs and seem to have no inner envelope."
At camp we made the following hasty notes: Vitellus 1.5, 1.5, 1.7, 1.6, 1.6, 1.4,

Rana virgatipes 445

1.5, 1.8, 1.6, 1.7, 1.8, 2.5 (embryo elongated) mm. ; envelope 5, 5, 5, 5.3, 4.9 x 5,
5, 6 X 6. 9, 5, 5.8, 5 X 5.8, 5 X 5.8, 6 mm. Occasionally jelly envelope flattened
spheroid-like. Jelly firm — often a flat surface where stuck to adjoining eggs.
Envelope single." "Envelope 5.2, 5.5, 5.2-5.8 mm.; vitellus 1.6, 1.5, 1.4 mm."
"Several fresh eggs have the envelope 3.8-4.2 mm."

The egg complement is rather small. One mass has 349 eggs. Another
had 474. "Some may reach 500 or 600 eggs but 474 was larger than some
complements. One mass looked to have little more than 200."

"The eggs are black above and a creamy white (not white as in Rana
sphenocephala and Rana pipiens) i.e. more creamy in fresh eggs than these two
species. Soon the black encroaches on the light vegetative pole. The eggs
look far apart as do Rana sylvatica eggs and being without an inner envelope
the parallel goes farther in that the outer envelope of Rana sylvatica gets to be
as much as 5.8 mm. as does the envelope in this species. Very fresh eggs
sometimes look to have sulphur yellow vegetative poles and black animal
poles." Comparisons with Ridgway colors made the colors "primrose yellow,
naphthalene yellow or light chalcedony yellow. Not so yellow as in Rana
palustris eggs."

The eggs and egg mass were identified presumptively. We neither saw
the eggs laid in the field nor did we capture mated pairs. They are Rana
virgatipes for these reasons :

1. The female taken the night of July 21 was near where a male Rana
virgatipes was croaking. We brought her into camp. On July 22 in the after-
noon Mrs. Wright when photographing the female accidentally pressed some
eggs out. Later July 23 (morning) the female though unattended by a male
had laid eggs in the jar. These checked with the eggs of the masses of the
field. This was in the nature of direct identification. No end of male Cope's
frog were croaking in the area where the female was taken. This is presump-
tive evidence.

2. They were not films as in Rana grylio and Rana clamitans. On July 23,
1922 we held there were only four anurans in Okefinokee swamp whose eggs
were not surely known : Scaphiopus holbrookU, Pseudacris nigrita, Rana aesopus,
and Rana virgatipes. They could not be the first two, no gopher frogs were
calling that night nor of the first two species. The presumption was that they
were R. virgatipes. The eggs were of the type of Rana sphenocephala or Rana
palustris but without the inner envelope. In this respect they were Hke R.
catesheiana eggs (but these are in films). No R. sphenocephala were calling
when we were at area the night of July 22. A mass of this species, however,
was later found.

3. On the morning of July 25, 1922, at the boat landing (strand or sphag-
num run) "Ridley and Mattie Chesser found a Cope's frog in the boat. It is
a female, gravid. Must have leaped into the boat and couldn't get out. This
female is smafler than any of the four males captured. When Anna (Mrs.
Wright) was cleaning her off for photography the female voided a few eggs.
Later I did the same and secured more eggs. These vitelli were somewhat less

446 Frogs of the Okefinokee Swamp

than those in the field, being 1.3-1.4 mm. and the inner envelope absent. The
jelly did not swell out quite as well as in the field. Our identification of the
egg masses of July 22 is certainly correct."

From subsequent studies of the eggs we found the jelly envelope to be
3.8-6.9 mm., the mode 5 mm. the average, 5.4 mm.; the vitellus to be 1.5-1.8
mm., mode 1.6 mm., average 1.6 mm. Egg complement 200-600.

Dangers. On July 22, 1922, when most of the masses were fresh we found
one mass about at the hatching stage. It must have been laid in the generally
rainy (July 18-21) period for southern Georgia. Doubtless it was laid near the
edge of the high water mark. By July 22 receding began. The mass was in
water only 3 inches deep and one half of the eggs were white and spoiled.
This is the only mass we found caught in this fashion.

HATCHING PERIOD

On July 22, 1922, the masses were all fresh except one. This was laid in
the rainy period from July 18- July 21. In the period from July 18-22 the
air minima were from 65-73°, maxima 84-96°. The water doubtless in this
shallow area kept about 70° or higher. These eggs were probably about 4-5
days in hatching. The egg masses of July 22 in the laboratory hatched in
about three days.

MATURE TADPOLE

Color description from life (May 23, 192 Jj). One tadpole very large,
brownish olive mummy brown or brownish olive. In general dorsum very
dark. A few large widely scattered black spots on the dorsum. The brownish
olive on rear sides surrounds lemon yellow or greenish yellow spots. This
yellowish color on either side of belly with a few small dark flecks. Middle of
belly pale grayish vinaceous or vinaceous buff. Ventral pectoral and branchial
region grayish violaceous blue or body color with vinaceous buff and greenish
yellow spots.

Tail. In general a grayish color. Along the middle of the upper crest is
row of large spots or these connected. This a much broader and more con-
spicuous row than R. grylio, being about 2 inches long. In middle of muscula-
ture is a dark fine or area to tip of tail parallehng the upper tail crest line of
black. The upper tail crest is irregularly black-margined to tail tip. This
black more or less outlines pale chalcedony yellow spots. The musculature
has numerous pale grayish vinaceous-vinaceous buff spots. Lower tail crest,
narrow with irregular black margin in tip region for i-i| inches. This crest
with numerous pale chalcedony yellow or vinaceous buff spots. Iris largely
black with ochraceous salmon or greenish yellow flecks.

One tadpole near transformation buffy citrine, olive lake, or dull citrine a
very few inconspicuous spots on dorsum. Tail crests with no black muscula-
ture with the vinaceous buff spots. Legs olive lake in color. Iris pinkish
vinaceous and black. Belly pale chalcedony yellow, or sulphur yellow.

Mouth parts. Teeth 2/3 or 1/3.

Plate XLIII
Carpenter frog ( Rana virgatipes)

1. Male, Chesser Id., Ga. July 22, 1922. Ventral aspect, x 1.2.

2. Female, Chesser Id., Ga. July 22, 1922. Ventral aspect, x 1.2.

3. Eggs in situ, second "Dreen," near Chesser Id., Ga. July 22, 1922. x i.o.

4. Rana grylio transformed for comparison with No. 5. Billy Id., Ga. April 26,

1 92 1. Ventral aspect, x 0.4.

5. Rana virgatipes, Chesser Id., Ga. July 15, 1921. x 0.33.

6. 7. Tadpoles, with two legs, with four legs. Lakehurst, N. J. July 4, July 7,

1923. X 1.0.

Rana virgaiipes 447

Measurements. Length of body (25-30 mm.) in tail (48-63 mm.) i. 89-2.1,
average 1.99. Width of body (14.5-20 mm.) in its own length i. 45-1. 86,
average 1.61. Depth of body (13-17 mm.) 1.1-1.2 in body width, average
1. 14. Depth of body 1.66-2.07 in body length, average 1.84. Depth of tail
(16-21 mm.) in length of tail 2.9-3.6 average 3.12. Muscular part (8-1 1 mm.)
1.62-2.0 in depth of tail, average 1.84. Spiracle 1.17-1.33 nearer base of hind
legs or vent region (14-17 mm.) than the tip of snout (17-20 mm.), average
1.23. Spiracle to eye (8-1 1 mm.) i. 42-1. 82 nearer eye than spiracle to vent
(13.5-20 mm.) average 1.62. Eye i. 0-1.22 nearer to tip of snout (7-9 mm.)
than eye to spiracle (8-1 1 mm.), average 1.16. Nostril i. 0-1.4 2 nearer eye
(4-4.5 mm.) than eye to snout (4.5-5.5 mm.), average 1.20. Mouth (5-5.5
mm.) usually i. 0-1.3 7 larger than internasal space (4-5 mm.), average 1.12.
Mouth 1. 3 6- 1. 8 in inter orbital space (7-9 mm.), average 1.57. Internasal
space (4.5-5 mm.) contained 1,55-1.87 in interorbital space, average 1.74.

The dimensions of the largest tadpole are :

Total length

92.0

Spiracle to vent

16.0

Body length

30.0

Spiracle to eye

II .0

Body depth

15-0

Eye to snout

9.0

Body width

18.0

Eye to nostril

4-5

Tail length

6s. 0

Nostril to snout

5-5

Tail depth

21 .0

Mouth

50

Musculature of tail

II. 0

Interorbital distance

9.0

Spiracle to snout

20.0

Internasal distance

5-0

General remarks. The eggs of July 22, 1922, hatched out in camp and we
kept the tadpoles for a month. The tadpoles a month old had grown very
little beyond the size of one week-olds. Our conditions were poor for them.

The last of June, 1923, Mrs. Wright and I were at Lakehurst, N. J., for a
very short period and secured no tadpoles. It was probably too late for tad-
poles. On May 22-23, 1924, "in a cut off pool from the main pond of Mr.
Emlie's pond where considerable sphagnum is are several tadpoles. We tried
to capture them with a dip net but didn't succeed. We resorted to a 10 ft.
minnow seine and caught 12 or 15 mature tadpoles. They must be Rana
virgaiipes or immense R. clamitans or Rana grylio. They cannot be the last,
yet in coloration have a black line on the upper and lower crests like the condi-
tion of Rana grylio. One tadpole had 4 labial rows of teeth. Associates of
these tadpoles are mud darters {Boleichthys fusiformis), mud sunfishes {Acan-
tharchus pomotis), little sunfishes {Enneacanthus ohesus), mud minnow (Um-
bra), American pickerel, and one adult greenfrog."

*Tn a left over ditch south of Noble's H. andersonii breeding pools found
plenty of Rana virgaiipes tadpoles, one with four legs and one transformed.
A little water snake in the ditch, also an eastern painted turtle. Mud minnows,
the commonest fish. In the large Emlie's pond heard an adult."

448 Frogs of the Okefinokee Swmnp

LARVAL PERIOD

In 192 1 we spent from April 26-Aug. i in the center of the swamp and
heard no R. virgatipes. In the last of July of that year we heard them on the
east edge of the swamp and at Black Jack Island in the south part of the
swamp. We therefore missed the tadpoles. In 1922 at Chesser Island we
found the species laying at July 22 but located no tadpoles from June 10-
Aug. 26. Why the mystery? In the first season we were where they were not.
In the second season we concluded they wintered over, transformed in May
and early June before we arrived and we were not there long enough after egg
laying for mature or half -grown tadpoles. Our trip to Lakehurst in 1923,
last of June, confirmed us in the supposition that they wintered over, and
transformed in May and June. We found only one debatable tadpole. We
therefore visited Lakehurst May 22-23, 1924, and found quite a few mature
tadpoles, transforming stages and two transformed individuals. We therefore
feel they winter over and this places them in the R. grylio, R. clamitans, R.
catesheiana, R. heckscheri, R. septentrionalis group in tadpole characters
though like others in egg-mass character.

TRANSFORMATION

Period. When we found transforming individuals on May 23, 1924, they
were scarce as were mature tadpoles. Stragglers transform to mid- July but
the vast bulk of wintering over tadpoles must transform in early spring by
May 15.

, Since this whole account was written we recently located two transformed
specimens of Rana virgatipes in our 1922 collections. We assumed we had
wholly missed this stage. These two transforming frogs were taken August
15, 1922.

Another addendum of June 8, 1929 is a record of one transformed frog
caught on lily pads in Lakehurst Lake by the Shinn brothers (Everett and
Whittemore) and R. D. Anderson. At the same time they caught four
adult males (51.5, 52.5, 54, 56 mm.) and a ripe female (56 mm.) in the same
places.

Size. On May 23, 1924, we secured from Mr. EmHe's pond 1/2 mile south
of Lakehurst two Rana virgatipes measuring 27 and 29 mm. respectively.
On June 5 some of the tadpoles of May 23 capture were approaching trans-
formation and the body length of these were from 25-30 mm. Another of
these tadpoles on July 14 transformed at 29 mm. and two on July 2 trans-
formed at 23 and 24 mm.

In 1923 (June 23) Mrs. 'Wright and I captured a 25 mm. specimen at
Lakehurst, N. J. Transformation may in rare cases extend to 32 or possibly
33 mm.

The two specimens taken in Okefinokee swamp August 15, 1922, are re-
spectively 28 and 3 1 mm. The first is completely transformed and the second
has a tail stub 10 mm. Each have the characteristic light stripe from eye to
above hind leg insertion as in adults.

Raiia virgatipes 449

We have, therefore, sizes from 23-31 mm.

General remarks. All the tadpoles taken at Lakehurst, X. J., May 22-23,
1924, were of the Rana damitans type. All of them seem to have the left arm
coming out first.

Transforming R. damitans from Ithaca and Adirondack Alts., N. Y., and
R. virgatipes from Lakehurst, N. J., compared are as follows:

Underparts and rear of hind legs and belly heavily blotched with black in
Rana virgatipes; finely dotted with black in Rana damitans. Light color of
under legs and belly to the level of fore legs, in fact to the chin, is white in
Rana damitans, in Rana virgatipes underparts to level of fore legs with some
vinaceous tint. Chin with few if any spots and greenish yellow or sulphur
yellow in Rana virgatipes: spotted in Rana damitans. Stripe on upper jaw
down to middle of side or farther sayal brown, clay color, or tawny olive or
buffy brown in Rana virgatipes; stripe in Rana damitans only to the shoulder,
lettuce green or calliste green on snout to light yellow green or sulphur yellow
near the shoulder.

GROWTH

C. S. Brimley (1909, p. 133) has supplied notes on the smallest specimens,
taken at Lake Ellis, Craven Co., N. C. ''Six specimens taken in 1906 and 38
in 1907, none being over 45 mm. in length of head and body. These little
frogs were found in shady places wherever there was water and in such situa-
tions acted much like cricket frogs, almost always coming to the surface im-
mediately after jumping into the water, and seldom diving to the bottom and
hiding there."

In the U. S. National Museum, Nos. 36629-31 were taken by Brimley
Bros. May 11, 1906, from Lake EUis and measure 32, 36, 42 mm. respectively.
The following May (24 and 25) 1907, J. Hurter secured Xos. 57675-78 from
the same place. They measure 43, 42, 38 and 39 mm. respectively. A speci-
men (U. S. Nat. Mus. No. 55447) secured at Lakehurst, N. J., by W. D.
Appel May 28, 1905, measure 35 mm. Two specimens (U. S. Nat. Mus. Nos.
37850-51) from the same place taken June 29, 1907, by W. T. Davis measure
40.5 mm. and 42 mm. The first specimens our collection ever received were
two taken by Mr. W. T. Davis at Lakehurst, N. J., May 27, 1906. They are
36 and 47 mm. — apparently two growth sizes.

My material falls into these groups: 23-30 mm. at transformation; 36-38
mm. one year old; 40-48 mm. 2 year old, and 49-55 3 j^ear old. The National
Museum material is from 32-39 mm. i year old; 40.5-47 mm. 2 year old, and
52-55 mm. 3 year old. Boulenger's (1920, p. 431) material falls into three
groups: 32-36 mm. i year old; 43 mm. 2 year old; 50-56 mm. 3 year old. In
general we believe they transform at 23-29 mm., or even at 31 mm., reach
32-39 mm. for i year olds, 40-48 mm. for 2 year olds, 49-56 mm. for 3 year
olds. The males begin to have enlarged thumbs and somewhat enlarged
vocal sacs at i year old. Might possibly breed at i| years old. Certainly
both males and females breed at 2 years and attain near maximum at 3 years.

Since the above was written I have examined the material in the Acad, of
Nat. Sciences, Phila. Cope's material of Oct. 1891 falls into sizes 31, 35, 49,

4SO Frogs of the Okefinokee Swamp

55, 55 mm. Early in the spring of 1905, April 23, T. D. Keim and H. W.
Fowler secured a large series at Great Egg Harbor River above May's Land-
ing, N, J. Twenty of them measure as follows: 39, 40, 42, 44, 44, 45, 45, 45,
45, 46, 48, 48, 49, 49, 49, 51, 51, 52, 52, 52, 53, 54, 55, 58 mm., the last two
females. Two specimens taken in summer on May 31, 1905 and June 20,
1901 were 45 and 50 mm. The fall (Oct. 189 1) group of Cope may show three
sizes: 31-35, 49, and 55; the early spring collection of Keim and Fowler
apparently fall into two groups: 39-49, 51-58. Or this material might be
interpreted as of two groups — the fall collection 31-35, 49-55; the spring
39-49, 51-58; the summer of two, 45 and 50. This would make half year olds
or less 31-35, year olds or less 39-49 and two year olds 49-58. Growth con-
ditions vary in N, J. and Ga. no doubt as does growth in bullfrogs in N. Y.
and La. (vide Viosca).

In the Museum of Comparative Zoology are the following:

3483 Lakehurst, N. J., W. T. Davis, 1905. 34 mm

2542 Lake Ellis, N. C, C. S. Brimley, May 24, 1907 37

2519 " " " " " " " " " " 36

2542 " " " " " " " " " " 34

3518 " " " " " " " " " " 42

712 Beaufort, N. C, J. G. Shute 1863 60
3560-68 Lakehurst, N. J., T. & F. K. Barbour, July 1914

35, 36 mm. past transformation

36.5, 40, 42 mm.

48 mm., 56, 66 mm.

Possibly these big representatives, 60 mm. and 66 mm. represent 4 year olds.

FOOD

Davis noted (1907, pp. 50, 51) that "These frogs domineer over one an-
other to some extent, and when insects were placed in the cage as food, it was
common for the more active individual, faihng in the attempt to catch a fly,
to turn on his companion and butt him until he retreated into the pool or
into a corner. The butted individual would hold his head down in the meekest
manner, and he became so cowed that if I touched him at any time with my
finger, he assumed the humble position. Miss Dickerson in The Frog Book
says that Rana pipiens and Rajia onca will snap at the head of a companion
frog that has taken a worm that he was trying to capture, but she thinks it is
probably not an exhibition of anger, but a desire to secure the disappearing
worm. However this may be, it is certain that the butting Rana virgatipes
in the above-mentioned case secured a great advantage over the other frog,
for after 'settling' his companion, he captured all of the insects."

ENEMIES

Every associate in the Okefinokee swamp and also at Lakehurst, N. J., if
large enough might feed on them. At Lakehurst, N. J., one of their worst
enemies is the water snake {Natrix sipedon sipedon). We saw several in the
sphagnum mat around the large lake. One specimen from Lakehurst had the
left arm from elbow down sharply cut off.

Rana virgatipes 451

AUTUMNAL DISAPPEARANCE

Published records and my own experiences carry their activities from
April 23-Aug. II. They come out earher than April 23 and must be active
at least two months after August 11.

AFFINITIES

(See R. grylio for comparison with that species).

A funny episode or joke on the author shows how closely Rana grylio and
Rana virgatipes parallel each other in coloration. Three days after we first
heard R. virgatipes in the prairie near Chesser Island we went (July 15, 192 1)
at night to a wooded causeway to catch some we heard. ''Harry Chesser
caught one. Another put into the water after he put his light on it. We
heard several. Caught two frogs which we mistook for Rana virgatipes until
on the next morning we examined them closely. They are young Rana
grylio. Two of the three real R. virgatipes we saw, escaped."

Boulenger (1920, p. 431) considers "The species does not exceed a length
of 60 millimeters from snout to vent, and may be regarded as a dwarfed form
derived from the R. catesbeiana type."

Previous to 1921, we knew the distribution of Rana grylio to be from
Florida and Georgia to Louisiana, that of Rana virgatipes from New Jersey
to North Carolina and that of Rana septentrionahs from Canada and Maine to
northern New York and possibly to Catskills. One might have thought the
three a series with a gap between each species. Now we know the first two
to overlap.

In coloration R. virgatipes looks most like R. grylio. In adult size it is
more like Rana septentrionalis but smaller or is possibly like a small R.
catesbeiana. At Lakehurst lake it reminds one of the water prairie form,
R. grylio; in the sphagnum strands and thickets of the Okefinokee it reminds
me of the peat lake species of Canada or beaver lake thicket form of Adiron-
dacks, namely Rana septentrionalis. In egg mass it is most like Rana septen-
trionalis, but in individual eggs unlike Rana septentrionalis, Rana clamitans
and Rana grylio it has no inner jelly envelope. The individual eggs are more
like the eggs of the film species Raria catesbeiana but the jelly is firmer. The
tadpole reminds one most of Rana grylio in coloration but it is more of the
Rana clamitans type. Like all of the Rana clamitans and Rana catesbeiana
type it winters over as a tadpole. It transforms at a size near that of RaTia
clamitans and may have a larger tadpole than that species. Therein it ap-
proaches R. catesbeiana, R. grylio and Rana septentrionalis.

Cope 1891 (pp. 1018, 1019) writes of the species as follows: "This frog is
not nearly related to any species of the genus. It has some points of re-
semblance to R. temporaria as the short posterior legs and moderate web:
but the interocular space is much narrower, the vomerine teeth more anteriorly
placed, and there are no dermal folds. In coloration there is no resemblance to
any other species." Of course Rana grylio was not then known. If it is to
have an European counterpart might it not be that Rana virgatipes is rather a

452 Frogs of the Okefinokee Sivamq)

dwarfed Eana escidenta or R. esculenta ridibunda. There might be those who
might derive our Atlantic coastal plain species all from Europe or place each
close to a European form e.g. Biifo foioieri and Bufo calamita, Scaphiopus
holhrooki and PeJohatcs, Eana virgatipes and Ratia esculenta, Hyla andersonii
and Hyla arhorea. There are difficulties in the way of such an interpretation
and some strong evidence for such an interpretation.

BIBLIOGRAPHY

1916 Barbour, T. Copeia, No. 26. January 24, 1916, pp. 5-6.
1919 Boulenger, G. A. Ann and Mag N. H. (9) III pp. 409, 413.

1919-1920 Boulenger, G. A. Proc. Am. Acad. Arts and Sciences, 55, pp. 415, 429-431.

1907 Brimlev, C. S. Journal Elisha Mitchell Soc. XXII, No. 2, p. 159.

1908 Brimley, C. S. Ibid XXIV No. i.

1909 Brimley, C. S. Proc. Biol. Soc. Washington, p. 133.

1891 Cope, E. D. American Naturalist, XXV, pp. 1017-1019.

1904, 1905, 1907 Davis, Wm. T. Am. Nat. XXXVIII, p. 893, 1905, XXXIX, pp. 795-
796; 1907, XLI, pp. 49-51.

1922 Davis, W. T. Jour. N. Y. Ento. Soc, Mar. 1922, Vol. XXX, No. i, p. 74.

1906 Dickerson, M. C. Frog Book, pp. 222-224.
191 8 Dunn, E. R. Copeia, p. 21.

1905 Fowler, H. W. Proc. Acad. Phila., LVII, 1905, p. 662.

1907 Fowler, H. W. Rept., N. J. S. Mus. 1906, pp. 120-126.

1908 Fowler, H. W. Rept. N. J. S. Museum 1907, p. 194.
1924 Myers, Geo. S. Copeia, p. 60.

1923 Noble, G. K. Zoologica, 1923, pp. 421, 422, 425.
1923 Pratt, H. S. Vertebrate Animals of the U. S. p. 183.

1917 Stejneger, L. G., and Barbour, T. Check List N. Amer. Amp. Rept., p. 39; 1923,
P- 37-

1906 Stone, W. S. Am. Naturalist, XL, p. 164.
1923 Wright, A.H. Copeia, p. 34.

1923 Wright, A. H. Bull. Ecol. Soc. Am., Vol. 4, No. 2, p. 9.

1923 Wright, A. H. Anat. Record 24, No. 6, p. 406.

1924 Wright, A. H. Proc. Biol. Soc. Wash., -^7, pp. 141-152.
1924 Wright, A. H. and A. A. Am. Nat., LVII, pp. 375-381.

Gastrophryne carolinensis (Holbrook)

(PI. I, Fig. 6; IV, Figs. 4, 5; V, Fig. 9; VI, Fig. 3; X, Fig. 14; XIV, Fig. 6; XV, Fig. 10;
XVII; XLIV; Text Figs, i, 24)

COMMON NAMES

Narrow Mouth Frog. Narrow Mouth Toad. Narrow-Mouthed Frog.
Narrow-Mouthed Toad. Nebulous Toad. Toothless Frog. "Rainy Day
Frog." "Rain Frog." "Frog-Toad." Carohna Tree Frog.

RANGE

Check list. "Virginia to Florida, Gulf States to Texas, Northward
through central valley to southern Indiana." — Stejneger & Barbour (1923,
p. 38).

Supplementary records. In Florida, Deckert (1922, p. 88) "Heard (Gastro-
phryne carolinensis) from a ditch beside the road. Royal Palm Hammock,"
Dade County. In Wilmington, N. C, G. S. Myers (1924, p. 60) took three.
Near Charleston, S. C, K. P. Schmidt (1924, p. 68) notes it at Mt. Pleasant.
In Alabama, H. P. Loding (1922, p. 21) records it from "Cherokee, Etowah,
Calhoun, St. Clair, Tuscaloosa, Mobile and Baldwin counties." In Alabama,
E. G. Holt (1922, p. 95) notes that "A specimen has been taken at Long-

Gastrophryne carolinensis 453

view, Shelby County, by L. S. Gibson, and I have collected the species at
Barachias, Montgomery County." In Doniphan County, Kansas, Jean M.
Linsdale (1927, p. 77) records "A narrow-mouthed toad was found on the
area in the summer of 1923." In Virginia (1918, p. 22) E. L. Dunn records
it in CaroHna County. The same year Philip H. Pope records it in Houston,
Texas. The next year (1919, p. 82) Doreen Potter records it in Lafayette
County, northern Mississippi.

These records of six years 1918-1924 represent its range rather roughly,
i.e., from Virginia, North Carohna, South Carolina, Florida to its tip, Ala-
bama, Houston, Texas, eastern Kansas.

Our own records for it in 19 17 begin just south of Richmond, Va., May 31.
We did not record it again until June 10 when we reached a low woods beyond
Tuskagee, Ala., and 30 miles from Montgomery, Ala. Our course through
Durham-Charlotte, N. C, Blacksburg-Anderson, S. C, Atlanta-LaGrange,
Ga., West Point-Tuskagee, Ala., was too far inland to record this Lower
Austral species. In 1926 (p. 83) we placed this species "Gastrophryne caro-
linensis Va.-Fla.-La.," as one of the nine Okefinokee species of the Lower
Austral frogs. Strecker (1909, p. 120) restricts it "principally to the humid
division of the Lower Austral Life Zone."

In 1922 on June 6 we heard it four miles north of Petersburg, Va., on June
7 at Cary, N. C, and on June 11 at Screven, Ga. (In connection with
Virginia records Dunn (19 18, p. 22) reports it from Carohne County north of
Richmond a considerable distance).

DeKay (1842, p. 65) writes "Dr. Holbrook thinks it possible that a species
may be found in this State, for he has heard its peculiar noise in the neighbor-
hood of New York; and Major LeConte informs me that he has seen a species
of Engystonia, said to have been found in a sandy district of this State." Sub-
sequent records have not yielded it in either New Jersey or New York and
fortunately DeKay placed it in the extra limital species.

No one has attempted a critical study of the Gastrophryne {Fngy stoma)
except J. K. Strecker in 1909 (pp. 115-120). His three conclusions were :

"First, that the narrow-mouthed toad usually reported from Texas is not
Engy stoma carolinense but Engy stoma texense which is a very distinct species.

"Second, that the Engy stoma carolinense Holbrook is exceedingly rare in
Texas. The only typical specimen examined was from Paris, Lamar County,
in the extreme northeastern section of the State.

"Third, that a small form with unusually pustulate upper surfaces,
peculiar coloration and short hind feet, from southeastern Texas, is worthy of
recognition as a distinct species {Engy stoma areolata Strecker)". . . In south-
western Texas the new species occurs in localities inhabited by the widely
"distributed Engy stoma texense, but in central Texas where the latter species is
the prevailing or, or as I am now fully satisfied, the only form of narrow-
mouthed toad, no examples have been found that are even an approach to
E. areolata.

"Engystoma carolinense Holbrook. This is the largest of the three forms
here considered and the most widely distributed. Examples have been ex-

454 Frogs of the Okefinokee Swamp

amined from Raleigh, North Carohna; Columbus, Georgia; Milton and Little
Sarasota Bay, Florida; Mobile, Ala.; St. Tammany County, La.; Hot Springs,
Arkansas; Cliff Cave and Butler County, Missouri; and Paris, Texas. It has
also been reported from Johnston and Wayne Counties, North Carolina,
Riceborough, Georgia; Clarcona, Lake Jesup and Micanopy, Florida; Green-
way, Arkansas; Calcasieu Parish, Louisiana; New Madrid, Missouri; and
various localities in Texas; but most of the Texan localities are open to doubt,
this species probably being confused with E. texense."

His conclusion (1908, p. 57) previous to his revision was that "The range
of this interesting batrachian was extensive, covering two-thirds of the State,"
but in 1909 he credits most of the Gastrophryne to G. texense, a view with which
we with less experience in the State feel inclined to concur.

Strecker (1915, pp. 46, 47) gives as its range in Texas "Eastern Texas south
to Victoria. Many of the published records for this species probably refer to
G. texense Girard, a smaller slenderer, paler, and more uniformly colored
animal with unspotted underparts. I have collected typical carolinense at
Cleveland, Liberty County, and have examined specimens collected at Paris
by Hurter, and Victoria by Mitchell. Miss Dickerson reports it from Hitch-
cock. Mr. Mitchell obtained only one specimen at Victoria, a locality in-
habited by both G. texense and G. areolata."

In 19 1 2 only five adults of the narrow-mouthed toad were secured, but of
larval stages to transformation a large series was taken.

GENERAL APPEARANCE

Holbrook (1842, Vol. V, p. 23) gives this "Description. The general form
of this animal approaches the oval; the skin is smooth, the head remarkably
small and short, though large for the genus; its extent is marked by a delicate
fold of the integuments behind the orbits; its shape is triangular, the snout
being very pointed; the upper jaw is dark brown, the lower dark grey; the
mouth is inferior and minute." Boulenger separates it from E. ustum (two
metatarsal tubercles) with a "single metatarsal tubercle." "Snout rather
obtuse, not twice as long as the diameter of the eye" separates it from E.
elegans, E. ovale and E. microps. H. Garman (1892, p. 7,^7,) calls it "a small,
clumsy toad, with a very small head and disproportionately stout hind
limbs."

C. S. Brimley (1907, pp. 157, 159) give its characters as "Upper jaw
without teeth. Skin smooth. Size small. Snout pointed. No parotoids ....
Hind feet not webbed." "Toothless Frog. Wake and Johnson Counties."
In 1926 (p. 81) the same author characterizes it as follows: "A transverse
fold of skin across neck just behind eyes, snout sharp, legs very short, un-
webbed. Size small, color dark."

COLORATION OF SPIRIT SPECIMENS (191 2)

In spirits the specimens are whitish or yellowish white below with a
prominent reticulation of brownish or grayish brown lines. Their color
scheme extends on the under side of the hind- and fore-limbs to the toes.

Gastrophryne carolinensis 455

Throat of the male slaty or blackish. Upper parts black or brown or gray.
In one specimen the whitish color of the under-parts extends on to the lower
tip and sides of the head as a grayish white on a dark background; the other
four have the head uniform black or brown.

STRUCTURAL CHARACTERS (WRITTEN IN 191 2)

Skin variable, smooth or finely tuberculate; head pointed, its tip obtuse;
upper jaw projecting beyond the lower jaw; nostrils much nearer the tip of
snout than the eye; a fold of skin extends from the insertion of one arm to
eye, thence to other arm insertion; tympanum concealed; toes free, subar-
ticular tubercle prominent; inner metatarsal tubercle present; in these five
specimens a depressed line across the breast from either arm insertion looks
hke an incipient fold. Tongue quite broad, ovate or elhptical, only the caudal
border free and this portion much thinner than the rest of the tongue; tongue
may be rounded or very shghtly emarginate. In none of the five specimens
is it so narrow as Cope figures and in only one so relatively long.

MEASUREMENTS

(1912-1914)

The five adults of 1912 measured 24.5-27 mm., average 25.8 mm.; the
head is 4.5-6 mm. long, average 5.0 mm., slightly less than the width of the
head at the rictus oris; the width of the head, 6-7.5 mm., average 6.6 mm.;
the head is contained in the length of the body, 4.1-5.2 times; snout, 3-4
mm., average 3.4, not twice the diameter of the eye, being usually i. 5-1. 2 5
times the eye (in agreement with Boulenger ('81, p. 162) and Cope ('89, p. 385)
but not with Dickerson ('07 p. 48) ; interorbital distance equal to the eye or
slightly more being from 2.25-3.5 mm. in width; eyehd 1.5 mm. or 1.6-2.3
in the interorbital distance; femur 9-10 mm., average 9.5 mm., equal to or
slightly less than the tibia which is 9.5-10.5 mm., average 9.9 mm.; tarsus
6-7.5 mm., average, 6.7 mm., about equal to the width of the head and
slightly more than its length; rest of foot 11. 5-12. 5 mm., average 12 mm.,
equal to fore-limb which is 11. 5- 12. 5 mm., average 12 mm,; forelimb slightly
longer than the distance from its insertion to the tip of the snout; posterior-
limb from groin, 27-32.25 mm., average 30.3 mm.; in none of the specimens
does the hind limb to heel reach beyond front of insertion of the fore-limb.
Miss Dickerson ('07 p. 48) gives ''head to shoulder one-third total length"
as distinctive of Gastrophryne texensis while for Gastrophryne carolinensis the
measurement is "head to shoulder one-fourth total length." In our specimens
this measurement is .4-. 3 3 of the total length.

(Recent Material)
Head to angle of mouth 1.4 (20 mm. c?')-i.44 (20 mm. 9 )-i-33 (28 mm.
cf )-i.43 (28 mm. 9 )-i.64 (36 mm. 9 ) in width of head; head to angle of
mouth 4.0-4.4-4.5-5.0-5.1 in length of body; head to rear of tympanum in
length of body; snout .83-.66-.50-.64-.60 in first finger;snout i.o-i.o-i.o-i.i-
1.1-.90 in fourth finger; snout 1.0-1.0-.70-.88-.70 in first toe; eye i. 5-1. 5-1.8-
1. 06-1. 6 in snout; eye 1.25-1.0-.90-.70-1.0 in first finger; internasal width

45 6 Fi'ogs of the Okefinokee Swamp

1.0-1.0-.72-.64-.66 in upper eyelid width; interorbital width .66-.75-.72-
.55-50 in upper eyehd width; interorbital width .66-75-1. 0-.87-.75 in
internasal width.

Forelimb: Forelimb 1.8-2. 1-2. 3 75-2.5-2.2 in length of body; forelimb
2.35-2.4-2.33-2.4-2.2 in hind limb; first finger 1.8-2. 5-2. 5-2. 5-2. 16 in third
finger; second finger i. 5-1. 66-1. 9-1. 5-1. 6 in third finger; second finger .83-
.66-.77-.58-.66 in first finger; third finger .88-.80-.92-.90-1.0 in second toe;
fourth finger i. 5-1. 66-1. 2 5-1. 46-1. 44 in third finger; fourth finger .83-.66-
.50-57-66 in first finger; internasal width 1.25-1.33-.90-.80-1.0 in first
finger; internasal width 1.5-2.0-1.1-1.35-1.33 in second finger; internasal
width 2.25-3.3-2.2-2.0-2.15 in third finger; internasal width i. 5-1. 5-1. 8-
1.3 5- 1. 5 in fourth finger.

Hindlimh: Length 1.3-1.17-1.0-.94-1.0 in hind limb; tibia 2.36-2.36-2.7-
2.8-2.5 in length; tibia 3.-2.76-2.7-2.64-2.5 in hind limb; tibia 1.3-1.1-1.15-
1.1-1.18 in forelimb; tibia 1,05-1,05-1.05-1.1-1.1 in hind foot without tarsus;
first toe 1. 33-1. 33-1. 64-1. 66-1. 8 in second toe; first toe 2.3-2.0-2.85-2.6-2,8
in third toe; first toe 3.0-3.16-3.9-3.66-4,0 in fourth toe; first toe 2,0-1.83-
2.5-2.33-2.2 in fifth toe; second toe 1.75-1.5-1.7-1.56-1.5 in third toe; second
toe 2.25-2.37-2.4-2.2-2.1 in fourth toe; second toe 1.5-1.37-1.52-1.4-1.2 in
fifth toe; third toe i. 28-1. 5-1. 375-1. 41-1. 4 in fourth toe; third toe .85-81-
.87-.90-.80 in fifth toe; fourth toe 1.0-.94-1.0-1.0-1.1 in hind foot without
tarsus; fourth toe .94-.90-.94-.91-1.0 in tibia; fourth toe 1.2-1,0-1,1-1,0-
1,18 in forelimb; fifth toe 1,5-1,7-1.57-1.57-1,75 in fourth toe; internasal
width 1. 5-2.0-1. 27-1. 07-1. 06 in first toe; internasal width 2. 0-2. 66-2. i-i. 7-
2.16 in second toe; internasal width 3.5-4.0-3.6-4.0-3.33 in third toe; inter-
nasal width 4.5-6.33-5.0-4.0-4.6 in fourth toe; internasal width 3.0-3.66-
3.1-2.5-2.66 in fifth toe.

HABITAT

Holbrook (1842, p. 24) writes "The Engystoma carolinense passes most of
its days in concealment, near old fences, or under the bark of fallen and
decaying trees, emerging only towards evening and after heavy rains. They
are frequently seen with myriads of the young of the Bufo lentiginosus,
apparently washed from their places of concealment by summer showers,
which has led many to suppose that they descended with the rain,"

"It is possible that Bosc referred to this animal when he says he observed
in Carolina, a 'crapaud bossu, ou une grenouille' living under the bark of
dead trees, though he describes its skin as so excessively delicate as to prevent
his preserving it alive even for a short time, in order to make a drawing of it.
Now, though the skin of our animal is smooth and delicate, I have kept
them alive for several months, and even sent them from Charleston to Phil-
adelphia, where they not only arrived in safety, but lived a considerable
time after,"

Le Conte (1855, pp, 430, 431) refers to Bosc as well. He writes "Very
common in the low country of Georgia under logs."

"In DeTerville's Natural History, Bosc confounds this animal with
Daudin's Bufo gibbosus. He says he found it in South CaroHna, but brought

Gasirophryne carolinensis 457

none thence to France, as the skin was so thin and tender, that he was unable
to preserve or even describe it; to me the skin appears as strong as that of
any other species of Ranina of the same size."

H. Garman (1892, p. 333) writes "It has been reported from the most
southern part of the State only, and is probably very rare even there. It is the
one species which, like the siren, water-moccasin and red-bellied horn snake,
mark southern Illinois as a part of a southern Zoological sub-region. Outside
of Illinois the species is almost confined to the Southern States. ... Le
Conte found it abundant under logs in Georgia, and others have collected it
among weeds. The pecuhar form, small immersed head, small withdrawn
eyes, and strong hind legs, suggest subterranean habits."

In 1894 Loennberg "found this peculiar little animal under old logs, dry
paLm-leaves, and such things, near lakes, and in moist places in Orange
County, for instance, in the pineland at Clarcona, and in the hammock
bordering Lake Jesup. It is not very common."

In 1893 Julius Hurter (p. 253) near St. Louis found "This little toad is
quite rare. I have only found it in spring at Cliff Cave, St. Louis County,
Mo., nearly on the top of a bluff on the southwestern exposure. They are
not easily seen on account of their color and being partially hid in the ground,
under rocks and logs." In 1897, the same author (1897, p. 503) writes of it
in Missouri. "Dr. Kennicott sent some that he caught in New Madrid
County, Mo., to the National Museum. I have found some specimens in
Butler County, and three at Cliff Cave, St. Louis County, which is, to all
appearance, the most northern locality of this subtropical species. These
toads are found under rocks, sometimes on the top of the bluffs. They are
very hard to see, as they are partly hidden in the ground and also protected
by their color."

In 1895 Rhoads (p. 396) says "The specimens taken (in Tennessee) were
found under logs in woodland near running water. Specimens: Raleigh
I ad.; near Chattanooga, i ad."

Percy Viosca, Jr. (1923, p. 40) in discussing the sea beaches of Louisiana
writes "Where the beach connects with some alluvial ridge, oaks, bushes
and other ridge land forms make up the vegetation and the fauna partakes
of the nature of the alluvial species Hyla squirella and Gastrophryne caro-
linensis sometimes reaching to the very edge of the Gulf." In (1926, p. 310)
he places it in the "Species common to the Atlantic, East Gulf, Mississippi
Alluvial and West Gulf Coastal Plains."

In 191 1 and 19 12 Deckert (1914, No, 3, p. 3) finds at Jacksonville, Florida,
"Engystoma carolinense . . . abundant everywhere after heavy rains." In Dade
Co., he in 1920 (192 1, p. 21) says "Of this, the 'Narrowmouth Toad' about
a dozen specimens were taken in Miami, on the borders of a rain-pool in an
empty lot at Miama Ave. and 22nd Street. Both sexes were represented. They
were hiding under pieces of limestone in black, mucky soil and were heard
calHng during the night of May 16 in company with Spadefoots and other

458 ^ Frogs of the Okefinokee Swamp

toads. Several were found at Lemon City under pieces of limestone in a
wagon track leading to a rock-pit, and one under a stump in a rock-pit at
Little River, in November."

In 19 1 2 we made a few notes on its habitat. They are: It frequents the
small ponds around the wooded edge of the islands where some shade is
afforded. One performer we took on some fallen saw-palmetto bushes a few
inches above a heavily shaded shallow pool which was 3 feet in diameter.
This pool was near our camp well. Another specimen was taken under dead
leaves. The species is not common but quite widespread. During our stay
we took three specimens: one on Honey Island, June i ; one on Billy's Island,
June 6; and another, same island, June 12. After July 15, 19 12 we received
two specimens one taken between July 15-Nov. 15, 19 12, and another between
Dec. 23, 1913-Aug. I, 1914. All in all, this species seems decidedly nocturnal,
a lover of rain, cover and moist situations.

(For 1 92 1 and 1922 records see Voice, Ovulation, etc.).

FIRST APPEARANCE

In 1 92 1 immediately after we entered the swamp on April 24 "one lad
found one under a decaying log." In Texas we found the related species
G. texense breeding as early as March 25. One can dig Gastrophryne texense
out of logs in Texas in February and doubtless the same can be done in the
southeast for G. carolinense. At Biloxi, Miss., G. S. Miller Jr. (U. S. N. M.
No. 5 1 120) secured the species in the middle of February.

GENERAL HABITS

Metachrosis: Le Conte (1855, p. 430) says it is capable of "Varying at
will very much in color, from dusky to brown, olive-cinerous or yellow cin-
erous, more or less varied, spotted or speckled with black, beneath pale brown-
ish white, punctate so as to appear spotted."

Variations in color: Holbrook (1862, pp. 23, 24) gives its general color
as chestnut brown above. Hallo well (1856, p. 252) has it olive. Gunther
(1858, p. 52) has one "rose-coloured." Dumeril et Bibron (1842, p. 743)
gives "brun olivatre ou marron." Boulenger has it (1882, p. 182) "brown."
Cope (1889, p. 386) considers it "chestnut."

H. Garman (1892, p. 332) discusses coloration and its variation at length:

"Color above olive-brown or gray, marked and spotted with dusky;
below pale yellowish, closely marbled with purplish, but more yellowish
posteriorly on the abdomen and under side of the femora. Two wide, poorly
defined pale bands begin at the fold of the skin behind the eyes and pass
backward and slightly downward to the insertion of the femora; they are
bordered above by a sinuous band of interrupted elongate dark spots, and
below by a wider continuous dark band, which in front passes immediately
over the fore legs, through the eye and around the snout, where it unites
with its fellow of the opposite side. Two dark bands cross the tibia.
The throat of adult males is bluish black. The colors vary with age and, to

Gastrophryne carolinensis 459

some extent also, at the will of the animal. Old examples are darker, and
the markings are in them more obscure. The characteristic markings are
consequently more apparent on medium sized specimens because of the
paler color and consequent greater contrast between it and the dark marks.
Examined with a lens, the skin of the body is seen to be sprinkled with minute
dark specks, the closer aggregations of which form the dark spots, while their
absence in numerous small irregular areas on the abdomens of the younger
examples produces a fine mottling of the under side. Occasionally the pale
bands on the sides of the back are so nearly the shade of the ground color as
not to be apparent; and they may be rendered still more obscure by the
absence of the dark band which generally bounds them above. A very young
specimen before me has a series of small dark spots along the middle of the
back. The feet are more or less spotted with dark above. A black spot over
the vent seems to be constant."

Strecker (1909, p. 117) notes that upper surfaces may be black, brown or
gray and under surfaces gray or light brown, speckled with white or light
yellow.

General habits: It is so secretive its habits have been little discussed and
most of these pertain to its voice. H. Garman (1892, p. 333) held that "Of the
habits but little can be written at present." Deckert (19 14, No. 9, p. i)
holds "the 'Narrow-mouth Toad' is a common Batrachian here (Jackson-
ville) but owing to its nocturnal habits, it is not often seen."

This is subterranean species. Its appearance alone indicates it. Never
during the day would one find it if he did not hear it bleat during a rain
storm or cloudy weather or if he did not unearth it from beneath cover.
Often in tearing decaying logs to pieces one finds them within the logs or
merely beneath them. Some boys found some under some hay cocks. Rarely
during showers one may chance on them by day. Even by day e.g., July 3,
192 1, one marvels at the numbers which may appear by day in a drenching
rain. Sometimes the din and population is incredible. The species appar-
ently does not have far to migrate at such times.

At times these creatures almost crawl along. They can hop from 2 to 6
or even 9 or 10 inches. Some think they can swim rapidly, others hold them
clumsy. At breeding times they often seek cover near the breeding place.
For example, on May 22, 1921, at Newt Pond we found eggs laid the night
before. In the daytime we turned over a piece of bark which was sticking
into the water. A Gastrophryne carolinensis male scurried or scrambled or
rapidly crawled into the water, but not too rapid for its capture. We have
noted under Voice how the males crawl out of the water up the vertical banks
at times. One male which the boys found under cut grass chmbed the vertical
side of a 2-quart can to the top.

VOICE

Holbrook (1842, Vol. V, p. 24) says 'Tt makes a feeble chirp at night
and at times when captured; and being but a clumsy swimmer, if thrown into
water it repeats this chirp in its endeavors to escape."

460 Frogs of the Okefinokee Swamp

In 1855 Le Conte (1855, p. 431) closes his catalogue with the following
note: "There is in the water of ponds and ditches a small frog whose note
exactly resembles the bleating of a lamb, so truly as to deceive any one.
This animal I have seen, but never succeeded in catching it. It is very
common in Georgia and I have heard it as far North as Norfolk in Virginia.
It is left to the inhabitants of those parts of our country to determine what
it is." It is unmistakably Gastrophryne carolinensis.

In 1891 J. A. Ryder (1891, p. 838) writes "This interesting Batrachian
occurs in the Piedmont section of North Carohna, near Littleton, and within
twelve miles of the southern boundary of Virginia. Its presence after a rain
may be discovered by its peculiarly plaintive note."

In 1914 Deckert (No. 9, p. i) writes "After thundershowers, however,
every pool and ditch resounds with its cry, which sounds Hke a nasal 'baa,'
or 'bee' and has also some resemblance to the noise made by an electric
buzzer." In 1924 (p. 60) G. S. Myers writes of it at Wilmington, N. C. as
follows: "The call, as heard in the terrarium reminds one of a loud electric
buzzer."

In 1 9 18 Philip H, Pope (19 19, p. 94) found them at Houston, Texas,
"June IS, 19 18. A heavy thunder shower brought out the narrow-mouthed
toads. Found them abundant and noisy in a large rain pool about ten o'clock
at night. The call is harsh and grating, almost a bleat. It is repeated fre-
quently but not as rapidly as that of the Hylas. The males sit in the water
in the thick grass that fringes the pools, and are almost impossible to find with
a light, although they are not at all shy. Collected six specimens by treading
down the grass into the water and catching them as they swam. They swim
slowly with their short legs and webless feet and do not dive when disturbed
as frogs and toads do. From this date until the middle of August I heard
them frequently, but seldom in any abundance. After a rain one could be
heard now and then in a pool or roadside ditch, before sunset as well as at
night.

In 1 91 2 we made the following notes: This species was seldom heard
during the party's stay within the swamp. Only on the 6th and 7th of June
did we hear it. In the evening of the 6th, this curious frog note arrested our
attention. To some it sounded like the bleat of a young lamb; to another
it resembled the note of a woodcock. All agreed it was very nasal and wholly
unlike that of a toad. As one approached the frogs the shrillness of the note
became very marked. On the following afternoon the frogs began again
during a rain. The natives associate it with such weather and term it the
"Rainy Day Frog."

In 192 1 our first records on calling came on May 21. Then and on the
day before the Gastrophryne began to "bleat like a lamb or calf. They
stopped to find Gastrophryne in a ditch. Sometimes they are under the
bank. One was thus. Another was out on the side of the bank. Its throat
like a Bufo terrestris throat. Black in color." On that date Mr. Harper
and I counted the intervals and periods of calls. His record of these 3 7 inter-
vals is from 3-58 seconds, at least 20 of which were 30 seconds or more and

Gasirophryne carolinensis

461

the average 30 seconds. The actual call in most instances occupied 1.5-2
seconds. The intervals of another calling frog were from 1-60, only one
below seconds, average 29 seconds. A third frog noted by Mr. Harper on
June 26 has 79 intervals from 3-59 seconds, average 30 seconds.

Normally one would not think of Gastrophryne's call as far reaching.
Frequently other frogs such as Hylids {Hyla femoralis, Hyla squirella, Acris
gnjllus), Scaphiopus or Bufos {Bufo querdcus and Bufo terrestris) may be in
chorus in the same place and may break the record or drown its calls. Some
of these species interfere with each other or Gasirophryne in other ways.
Once in a flooded furrow, July 3, 192 1, we saw two mated pairs of Gasiro-
phryne. One we captured but before we could try for the other a {Bufo
terrestris) toad had butted the pair and frightened it. To return to its voice.
One has to be within 20-60 yards to hear Gastrophryne calls under normal
conditions. They usually call from the water with rear parts submerged
and only fore feet planted on back or some support. Occasionally they may
be amongst the trash on surface. Occasionally they are out of water on the
bank or in a grassy tussock. On May 26 we observed numerous males in
one ditch. At times they would crawl out of the water and up the vertical
banks of the ditch a few inches. The white throat is swelled out Hke the
bubble of a toad or spadefoot and the frog becomes vertical or bent backward
with the effort. In the interval the frog returns to a position short of vertical,
deflates the throat but may pulsate the throat. At times the male has one
or two other notes not of the b a a a . . . order. The same is true of Gastro-
phryne texense.

1921

A number heard. Air 68°.

A chorus in the evening. Air 70°.

Commonly heard in the evening, also in mid afternoon.

A few caUing tonight. Air 77°, 10:09 P- ni-

At 12:30 a. m. a great chorus. More Gastrophryne calling than ever

before. Air 73°.

Rain of two inches. More Gastrophryne than any other time. No

end of Gastrophryne.

Several in late afternoon. Air 71°.

Tonight after 2 inch rain all along ditches and elsewhere heard.

Abundantly caUing at night.

Heard plenty of Gastrophryne.

Common tonight.

Calling in evening in chorus.

Not so common this evening.

One calling near cypress bay.

Heard 5-6:30 p. m. after rain. Common in evening.

Less heard than formerly this night.

Flood of rain. Never heard such a din.

A few calling. Air 75°.

6. A Gastrophryne or so; p. m. rain. Several calling in the evening.

7. Commonly heard.

15. One heard.

16. One heard about 7:15 p. m.
23. Gas/rop/irT/we calling tonight.

May

June

June

July

462

July

26

"

29

"

30

Aug.

5

"

7

"

16

Fi'ogs of the Okejinokee Swamp

Last night many heard.

One or two heard.

No end of Gastrophryne calling in the evening.

Heard.

Heard.

Several heard.

July

Aug.

1922

Heard a few between 8:30 p. m. and midnight.

Common at 7 p. m. in meadow below spring.

Downpour. Heard a few in evening.

Heard near spring in evening.

Heard quite a few narrow-mouths 8-1 1 p. m. after storm.

Common at night along Dixie Highway.

Common.

Commonly calling at 10 p. m.

Heard a few 8-1 1 p. m.

Several heard 10-12 p. m.

A few only.

Several heard.

Several heard at night.

Taking the voice record for 192 1 we find for stations around the swamp
the following data from May 20- July 29. For the day before the record
from May 20- June 5 the maxima have an average of 84.5° and range from
78-94° the minima, an average of 68° and a range of 63-70°. The period
from June 23 -July 29 have maxima average 92.5° range 80-97° and minima
from 67-78°, average 71°. Or the whole period from May 20- July 29 have
air maxima of 78-97°, average 89°, minima of 63-78°, average 78.5°.

For the day of the record air maxima from May 20- June 5 are 78-94°,
average 85,5°, minima from 62-70°, average 68°. From June 23- July 29
the maxima are 80-97°, average 91.5°, the minima from 67-74°, average 70°.
The whole period May 20- July 29 give air maxima from 78-97°, average 89°
and minima from 62-74°, average 69°. These records for the day before
and for the day of the record correspond closely. The minimum range of
62-78°, averages 68°, 68°, 68.5°, 70° are more important than maxima but
not of the consequence of humidity.

Humidity is the important factor in this species in particular. They are
truly a "Rainy Day Frog." At three or four stations around the swamp we
have this record :

May

June

20

. 58 mches an

d I. 22 the da

21

•15 "

' -35 " "

22

.11 " '

' .56" "

24

.01 " '

26

.40 "

4

.67 " '

' .76 " "

5

2.56 ;; ;

' -39 " "

23

' I. 41 "

24

.98 " '

25

.70 "

' .98 " "

Plate XLIV
Narrow-mouthed toad (Gastrophryne cawlinensis)

1. Pool along the track, Billy Id., Ga. May 22, 192 1.

2. Male and female (larger), Billy Id., Ga. May 23, 192 1. Lateral aspect, x 0.75.

3. Female (larger) and male, Billy Id., Ga. May 23, 1921. Ventral aspect, x 0.75.

4. Egg clusters on the surface among driftwood, Billy Id., Ga. May 22, 1921.

5. 6. 7. Tadpoles, Trader's Hill, Ga. June 24, 1922. x i.o.

8. Transformed frog, Chesser Id., Ga. June 29, 1922. x i.o.

9. 10. II. Males croaking, Billy Id., Ga. August 4, 1922. Flashlights.
12. Eggs, stained, Chesser Id., Ga. July 4. 1922. x 1.3.

-7' ■'- S 9

6 ^ , '

>^R^ • • • •' * •

12

Gasirophryne carolinensis

463

June 26 2 . 59 inches and .70 the day before.

28 .57

" 29 3.02

" 30 2.08

July I T

2 .68

3 3-47
" 4 I. 00

6 4.56

7 .89
" 26 1.32
" 29 2.84

(For temperature of chorus records see Ovulation).

::

' -57 " '

' 3-02 " '

' 2.08 " '

' T " '

' .68 " '

\ ^'^'' "„ \

' 4.56 " '

' 1.70 "

' 5-13 "

MATING

Male {From life, April 25, 1921) (Non Ridgway). Top of head and upper
eyelids mineral gray, pale smoke gray or with yellowish cast. June 5, 192 1.
Under parts dusky and bluish- white. White on throat, fine dots not blotches;
therefore throat looks blacker in male. All over back dusky with fine bluish-
white spots or dots. Pupil circular. Iris black with bluish-white.

Female {From life, June 9, 1921). Lighter, larger. Practically no spots
on pectoral region. None on throat. Throat same color as belly. Practically
no rusty spots. Ground color of the dorsum more greenish, i.e., olive gray
or mineral gray or grayish olive. Sometimes at breeding season males and
females may be alike in color, e.g., on April 24, 1921, several were thus, several
pairs reddish, one pair gray. Most of pairs however were diverse.

Structural differences. From its first description authors have noted males
with a sub-gular vocal vesicle." Or put in the negative H. Garman (1892,
p. 330) has males with "no saccular dilations of the skin at the angles of the
mouth." His affirmative is "Males with an internal, sub-gular vocal sac."

We found that in the Okefinokee frogs there were some males revealed
externally at 21 mm. and females at 22 mm., and our largest male was 29
mm. and largest female 30 mm. In the whole range of the species we found
males and females so indicated externally at 20 mm., the largest male 30
mm. and the largest females 31-36 mm. They apparently begin breeding
the second spring or summer after transformation.

The male's throat in alcohol may look black, bluish-black, bluish-gray,
or in many cases darker than the rest of the underparts. The female has the
throat much as the other underparts though the light or bluish-white spots
on the throat are smaller than on the venter proper. The male's throat
seldom has the lighter spots. Frequently the males in alcohol have the lower
throat just ahead of pectoral region thrown into one or two transverse folds
or plaits, sometimes it appears as one or more ridges. The lower jaw of the
male often has light spots and is brownish-black like venter, but often it is
uniform self color like the throat.

The actual measurements of several mated pairs show the males usually
smaller. For example one ovulating pair of July 4, 1922, caught at midnight

464 Frogs of the Okefinokee Swamp

were 24 mm. for the male and 28 mm. for the female. Another ovulating
pair of the year before, July 3, 1921, were 23 mm. cf and 27 mm. 9 .

Duration, night or day. Amplexation. This subterranean species normally
breeds and mates at night. Heavy, warm rains of 1-4 inches may precipitate
mating in dayhght and rarely one captures a croaking male in mid-day.
We took or recorded mated pairs May 21, July 3, July 7, 192 1, all taken at
night choruses. In every one the amplexation was axillary except in one
pair where for a time it was preaxillary.

OVULATION

Habitat. Deckert (1914, No. 9, p. i) has "caught hundreds of these
queer httle toads, and on August 28, 192 1, came across their spawn in a ditch
between two potato hills."

In 192 1 we found egg masses or mating or ovulation in the following
places: The clearing on Billy's Island was in a hammock with pine barrens
on the south and cypress bay on north end. Between the lumber quarters
and the bay were the Lee's cleared fields. In the temporary pools in the
streets of the lumber quarters after a rain or in any depression they might
lay. In the furrows of corn fields or in a walled-in area in a pumpkin patch
or in a well beside a shanty they laid. In small ponds at the edge of the
clearing or edge of hammock or edge of bay they frequently deposited eggs.
The most prevalent places were the ditches in the east and west negro quarters
beside the railroad tracks. In any place where a depression will furnish
water they ovulate, — places such as a sink hole near a stable, temporary
pools in old camp grounds (191 2 camp site), pools around a landing, overflows
or depressions around a turpentine still or building. At times after a 2-4
inch rain they may appear all over the island in any habitat with water.

In 1922 we found them in deep dug pools on high banks of St. Mary's
River at Camp Pinckney, in pools beside wood roads, in ditches along Dixie
Highway, in open, artificial and natural ponds in cut over pine barrens, in
ponds of Ilex myrtifolia or Hypericum fasciculatum or shallow cypress ponds
or in moist pine barren pools normally dry or in wet meadows.

Apparently whenever a great rain comes they breed wherever much
surface water accumulates. Apparently they do not lay so much in the
deeper pools and in this respect are much hke the fast developing Scaphiopus
holhrookii.

Period. J. A. Ryder (1891, p. 838) holds that "Oviposition seems to
occur in the evening and during cloudy afternoons. It is now late in July,
yet two lots of ova have been found by me which have been very recently
deposited by the parent female 'frog-toad' as it is known here amongst the
natives."

In Raleigh, C. S. Brimley (1896, p. 501) writes that "This species is very
abundant in the breeding season, which is in July and August, and possibly
the two preceding months. Have never seen any except when breeding; I
think they are nocturnal." And Strecker (1909, p. 120) reports that C. S.
Brimley says that it "breeds from May to August" at Raleigh, N. C.

Gastrophri/ne carolinensis 465

In 192 1 "we did not find the eggs of this species but the 'pipers' of June
6 and 7 might have been tardy breeders."

In 1921 we have ovulation records May 20, May 21, May 22, May 26,
May 28, June 4, June 5, June 23, June 25, July i, July 2, July 3, July 6,
July 7, July 23. In 1922 we have such records on June 22, July 4, August
7, August 9, August 17. We have records of spent females May 23, June 8,
July 3, July 4, August 23, and of ripe or gravid females March, May 23,
June II, June 18, July 3, July 22, August, August 23. This gives breeding
May 23-August 23 or later.

In 1920 (pp. 29, 30) we wrote "The narrow-mouthed toad is a form whose
life-history is not wholly understood .... The author's limited experience
with the species suggests that the eggs are usually laid during the spring or
early summer." In February, 1923, (p. 34) we state that "From May 15
on to June ist, five more start (to ovulate)." Gastrophryne carolinensis is
the third in the list. In 1924 (p. 378) we give the "Season, May 21 to August
17." The records show May, June, July and August the normal months
just as Brimley long ago said. Doubtless they begin earlier if our March
breeding (1925) date for G. texense be indicative.

Temperature and humidity. When eggs were found the minima for the
day of the record varied as follows: 63-68°, average 66°, 64-70°, average
69.5°, 70-72°, average 71°, 69-74°, average 72° in 192 1 or 64-70°, average
67°, 70-72°, average 71°, and 73-74°, average 73° in 1922. These records
63-74° or averages 66-72° fairly represent the minimum air temperatures
of ovulation. The data for mated pairs are much the same. The optimum
temperatures are nearer the records for chorus which follow.

The air temperatures which we recorded at camp at the time of 1 2 choruses
of Gastrophryne in 1921 range from 70-77°, mode 70°, average 72.5°; the
records of 7 fair callings range from 68-75°, average 72.5°. In 1922 for 5
choruses we have 70-78°, average 74°, for four voice records not chorus
70-77°, average 73°. Like Scaphiopus it likes drenching rains but will come
out on less moisture than the former.

When these choruses, matings and ovulations came a heavy rain preceded
or immediately followed the activities. Usually 63-74° are frequent minimal
temperatues of ovulation but 70-77° seems the optima of minima.

EGGS

Attachment, egg mass, egg description. J. A. Ryder (1891, p. 838) is the
first to describe the eggs of this species. His description follows: "The
eggs are heavily pigmented at the upper or' animal pole, being darker than
the eggs of Rana, and also considerably smaller. They are laid in strings,
but so coiled as to form a nearly complete single layer over a considerable
surface of water. The gelatinous coating, as in Rana, spreads out under
the surface of the water, where by its adhesion to the layer of molecules at
the surface a certain amount of support is thus gained for the eggs. It is
therefore evident that surface tension is an important agent in keeping the
eggs of this genus, and those of Rana, at the surface of the water. Other

466 Frogs of the Okefinokee Swamp

genera, such as Bufo, do not have the eggs supported on the surface, but
are laid in strings formed of one row of eggs, wrapped in a gelatinous cord
which lies on the bottom of the pond in which oviposition occurs. Still other
forms have the eggs glued together in large masses and supported upon water
weeds; this is notably the case with some Urodeles, such as Amblystoma."

Deckert (19 14, No. 9, p. i) describes them as follows: "The eggs are laid
in oblong jelly-like sheets or flat masses, about i 1/2 inch long and i inch
wide. The egg-masses contain about 100 to 150 eggs."

In 192 1 we found them commonly. A few field notes from our journal
will reveal some of their characters: "May 21. Found eggs fastened to
vegetation beneath water some 1-2 inches, also saw some floating free and
some floating in a mass around a stick on the surface. They were black and
white. Thought them at first possibly Hjjla cinerea. Examination in the
camp reveals they have no inner envelope and they look like Rana clamitans.
Have the habit of free floating eggs. Last night and today it rained an inch
or two, so some eggs are immersed."

"May 22. In ditches by negro quarters were Gastrophrijne eggs. Each
egg stands out distinctly. Ditch 2 feet wide and 1/2-1 1/2 feet deep. Black-
ish, trashy, oily water. Packets of eggs floating. Some masses i x i inch
in diameter; others 2x1 inches; others 2x2 inches. Some round masses,
others square. Each jelly envelope abuts that of the next in pentagonal or
hexagonal fashion. In proper light a mass of eggs makes a mosaic. Some
eggs along banks in weeds may be i foot 10 inches long and 3 or 4 inches
wide. One mass in middle of ditch was amongst chips and was 4x7 inches.
Few such in mid pond. Believe they lay large masses along edges or amongst
brush. Wind scatters them and carries them around on water's surface."

"In a clean pond (20x3 feet) along its edges in amongst grass, one packet
of eggs, 100-125 floating. They are brown and yellowish. One mass 6 inches
long X 3 wide separated from another mass, round, 2I x 2I inches. These
three masses are floating films. More eggs in a mass than in a film of Hyla
versicolor. Insects get into the fresh masses amongst roots.

"We went to Old Hog's Hole. The eggs there we thought R. dimatans
are Gastrophryne. Whole pond covered with little masses. These must float
away from main mass. Over in Newt Pond amongst weeds are fresh Gastro-
phryne eggs. . . . Each egg stands out like glass marbles but attached to
each other. Believe eggs are laid in 2-4 large masses or one in amongst
brush, grassy edges. Then wind drifts them away in small floats. Finest
appearing frog's eggs which float in water. Masses may be at times even
triangular, one 7x7x7 inches. '

"May 26. As with other floating egg films, more Gastrophryne eggs in Old
Hog's Hole Pond are revealed by air bubbles amongst the jelly.

"May 28. New Hog's Hole Pond has many Gastrophryne eggs hatching.
Newt Pond has numerous masses of fresh and hatching Gastrophryne eggs
floating. In certain lights each egg looks like a circular oil spot on water.
52, 9, 48 are counts of eggs in three films. Sometimes in a shady place each
egg looks to be a milk-white circular ring with a clear hyaline center when

Gastrophryne carolinensis 467

hatching. One mass near bank 4x8 inches. When I approached in the
water the waves separated this mass into 4 masses of 74, 71, 86 and 85 eggs."

In 1923 (p. 34) we write that "Gastrophryne carolinensis spawns clear
marble-Mke eggs in films on the surface, these eggs truncated on the top."
In 1923 (p. 406) we hold that normally Gastrophryne carolinensis "Eggs (are)
laid in films." Of this species in 1924 (p. 377) we say that "One form, Gastro-
phryne carolinensis, although it lays at the surface, has the most beautifully
distinct, firm eggs of all the species considered." "Egg mass, a surface film.
Egg envelope outline always distinct, never lost in the mass; eggs firm and
distinct like glass marbles, making a fine mosaic ; envelope a truncated sphere,
the flat surface above; envelope single 2.8 to 4.0 mm.; vitellus i. 0-1.2 mm.;
color black above and white below. Egg complement 869." This egg com-
plement comes from the count of one of the 192 1 females. The truncated
envelope did not appear until Mrs. Wright tried to manipulate the eggs for
drawing. They were hard to turn over.

The detailed study of several eggs by Mrs. Wright gave no inner envelope.
The outer envelope ranged from 2.8-4.0 mm. Of 31 eggs, 3 were at 2.8 mm.,
II at 3.0 mm., i at 3.2 mm., 4 at 3.4 mm., 3 at 3.6 mm., 4 at 3.8 mm., and 5
at 4.0, average, 3.35 mm., mode 3.0 mm. Of 34 vitelli 17 were at i.o mm.,
2 at 1. 1 mm., and 15 at 1.2 mm., or average of i.i mm. We fear that our
field record of some eggs brown above and yellow below are of H. femoralis.
Do.ubtless it is a mistaken identification when we little knew this form's eggs.

Dangers. No species lays in more transient places than Gastrophryne
unless it be Bufo quercicus. W^hen one finds mated pairs hopping around in
sweet potato furrows or finds a congress in a wet meadow which will be
dry a few days later, he marvels that any of these forms come through to
maturity. Seemingly these subterranean forms breed right where their
retreat happens to be. Like Scaphiopus they wait for a drenching rain to
flood their locality. The toll from these transient breedings must be im-
mense. But even in ponds and bigger ditches great losses happen. For
example, on June 25, 192 1, we note: "Along the deeper ditches of west
negro quarters are plenty of Gastrophryne eggs. Many of these are getting
stranded even soon after laying. What a frightful waste there is in nature!"
Several times we found spoiled white eggs where the eggs were caught and
fastened to vegetation. Normally these eggs are free on the surface and drop
with the level of the surface.

HATCHING PERIOD

J. A. Ryder's (1891, pp. 838-840) developmental story might be apropos
at this point: "The development of the eggs of Engy stoma is rapid; three
days after deposit the larvae escape from the egg-envelopes. Throughout
the course of development there is well-marked evidence of geotropism, or
of the action of gravity in maintaining the equihbrium of the egg. The
animal or black pole remains uppermost, the heavier or light-colored vege-
tative pole remains lowermost; the whole egg is thus maintained in a position
of static equilibrium with the earth's center. There seems to be no tendency

468 Frogs of the Okefiiiokee Swamp

to rotate the egg through cUiary action, previous to the closure of the medul-
lary folds. That cilia are entirely absent on the eggs of Engysloma is proved
by the fact that at the time the medullary groove is still open every egg of
the same age is in exactly the same position in respect to the center of the
earth, and remains so for a long time, or until the tail fold is well developed
and the medullary groove has been closed. Before the closure of the medul-
lary groove, but after the egg has begun to elongate and the paired, secretory,
adhesive surfaces of the under side of the head have appeared as rudiments,
the position of the animal and vegetative poles is still the same as in the
undeveloped ovum. The head end of the egg is slightly elevated above the
caudal end. This is due to the forward growth of the head, and the retention
of the heavier yolk, farther backward under the posterior half of the medul •
lary groove. The medullary groove thus, comes to be inclined downward
a few degrees from the head toward the tail, but the groove looks exactly
toward the zenith, while the yolk looks downward in every egg, even the
inclination of the medullary groove with respect to the horizon being the
same for every egg of the same age.

"At this stage I neglected to note an extremely important fact,— Viz.,
whether the cephalic and caudal poles of the same row of eggs were all of
them lying in the same direction. The fact that no change of position occurs
for a long time in the eggs of Engysloma would indicate that possibly we
might find that the future cephalic pole of the egg bore a constant relation
to the cephalic pole of the parent Engysloma, such as is known to be the
case in Balrachus lau. Such relations between parent and offspring exist to
a marked degree, if they are not universal, in plants, and it is desirable to
know to what extent the same rule holds with respect to animals. According
to what has preceded, the early development of Engysloma is peculiarly
favorable for the purpose of testing the theory that the cephalic and caudal
polarities of the parent are transmitted directly to the offspring, or that the
future long axis of the embryo already conforms, even in the egg, to that
of the parent.

"The next step in the development of Engysloma is somewhat similar to
that of Rana. As soon as the larvae have the tail-fold well developed they
turn over and lie on the side, curved upon themselves, within the egg-evelope.
This is the condition of the eggs on the second and early part of the third
day. On the third day the larvae leave the egg, and then tend to fall upon
the bottom of the pool or receptacle in which hatching occurs. Soon after
this they begin to swim about actively, and, singularly enough, instead of
swimming like a fish, for some reason, which it is difficult to make out, the
larvae revolve on their own axes. This singular mode of locomotion is
probably due to the peculiar manner in which the tail is vibrated. This
mode of swimming lasts about a day, after which the larvae begin to swim
in the usual fish-like way. At this stage, when the larvae come to the surface,
the head is in contact with the surface of the water, and when quiescent the
axes of the body, when in a condition of equilibrium, assumes an angle of
about eighty degrees with the surface. The adhesive organs near the mouth

Gastrophryne carolinensis 469

now become functional. Up to this time the Ught area on the yolk is prom-
inent, and enables one to watch the singular rotation of the larvae. The head
now begins to widen rapidly, and the light area on the belly become darker.
The tail-fold soon becomes very thin, and bordered all around by a delicate
edging of black pigment. The larvae cease to rotate on the fourth day, and
no longer take up their angular position at the surface of the water, and now
behave very much like the larvae of Rana. They are now very heavily
pigmented over the whole of the bodj'^ ; the light area over the space where the
yolk was formerly placed has disappeared, and the larvae are now black as
seen from above."

Deckert's eggs found August 28, 191 1, but laid night of August 27, 191 1,
hatched on August 30. In 192 1 on May 24 I have the note. "Gastrophryne
eggs probably laid night of May 20-21, probably hatched night of May
23-24" or three days later. On the night of May 26-27, 192 1, when the
air was 72° a great chorus was on, and the following morning many places
had fresh eggs. On May 28 in several places some of these were hatching.
This makes i 1/2 days for hatching.

MATURE TADPOLE

Color description from life. General color black overlaid with very fine
light purpUsh gray or quaker drab or hair-brown dots. Transverse stripe
of belly divided in middle and is apricot yellow or buff yellow color. Along
either side of the belly a light buff stripe. Another such stripe on either side
of the gill region. Sometimes whole venter with small light buff or pale orange
yellow spot heaviest on belly and sides and hghtest on throat. The inter-
spaces are purpUsh gray, violet gray or plumbeous.

Tail with light buff or white stripe along middle of muscular part of tail.
After the first one-half inch the stripe breaks up into spots, which finally
disappear caudally. Above and below this continuous stripe is clear black.
Above this black is Hght purphsh gray or quaker drab dots. Quaker drab
not in lower crest. Lower and upper crests on caudal half with muscular
part heavily blotched with black or rather fight purphsh gray or hair brown
giving the tail tip almost a black appearance.

General appearance. Tadpole small, (26.4 mm.) flat, wide, elliptical,
snout sometimes somewhat truncate. Tail medium, obtuse or rounded,
sometimes with black tip. Dorsal or ventral crest not equal to to depth
of the musculature. The dorsal crest scarcely extends on to the body, reaching
a vertical somewhat ahead of developing hind legs. Spiracle medium, closely
associated with anus, just ahead of it, not very apparent until hind legs begin
to appear, when it becomes separated from the anus. Eye is on the lateral axis,
distinctly lateral in position. From one eye to snout and around to the other
eye the tadpole has a prominent canthus made by the flat ventral and dorsal
sides of the head. Anus median at the end of the edge of the ventral crest.
Muciferous crypts indistinct.

470 Frogs of the Okefinokee Swamp

Mouth parts: Teeth o/o. • No horny mandibles; no labial teeth; no papillae;
upper labial edge dark and quite emarginate in the middle; just below this
emargination is a lower light-colored median beak-like prolongation on the
margin of the lower labium.

Measurements. Length of body (9.2-10.8 mm.) in tail (9.8-16.4 mm.)
1. 06-1. 77, average 1.37. Width (6.5-8.0 mm.) of body in its own length
1. 3-1. 65, average 1.43. Depth (4.4-5.0 mm.) of body i. 35-1. 65 in body
width, average 1.5. Depth of body 2.0-2.3 in body length, average 2.15.
Depth (3.2-5.0 mm.) of tail in length of tail 2.4-4.25, average 3.15. Muscular
part (2.4-3.6 mm.) 1.16-1.66 in depth of tail, average 1.32. Spiracle just in
front of vent. Nostril within lateral edge of mouth fold. Mouth (2.0-3.0
mm.) contained 1.5-2.25, (average 1.93) in interorbital distance (4.5-6.5
mm.).

The dimensions of the largest tadpole are :

mm. mm.

Total length

26.4

Spiracle to vent

Body length

10. 0

Spiracle to eye

Body depth

4.8

Eye to snout

4-4

Body width

7.0

Eye to nostril

Tail length

16.4

Nostril to snout

Tail depth

5 0

Mouth

2.4

Musculature of tail

30

Interorbital distance

4.8

Spiracle to snout

Internasal distance

General remarks. In 191 2 we made the following extended notes on the
tadpoles of this species: "In a small pond at the edge of the Lee's clearing
and just outside the swampy wooded edge of Billy's Lake we found on June
12, 1912, the tadpoles of the narrow-mouthed toad. In the same pond were
Gamhusia affinis, Biifo tadpoles, Rana sphenocephala tadpoles and green
frog eggs and adults. The pond was 15 x 10 feet in dimensions and 3 or 4
feet deep. Here we took half grown, mature and transforming tadpoles of
this species. On June 24 the same stages could still be found as on the last
days of our trip (i.e., in July). Some of the tadpoles would not have trans-
formed before September or October. The description of the tadpoles
follows :

"The back is fuscous, olive brown or light brownish olive or raw umber,
rarely with a few white spots; the belly may be clear white. Oh the throat
the fine thickly set dusky specks of dorsal color surround small white areas;
the rest of the venter is mottled, blotched or reticulated, the white patches
being defined by a reticulum of dorsal color. The mottling is most prominent
on the sides of the belly. In some the belly is entirely white with no reticu-
lations and with fine speckings. In many on either side where dorsum meets
venter there is discontinuous white band from in front of the insertion of the
hindhmb well forwards toward the future position of the arm. The belly
coloration foreshadows that of the adult only in the latter the white spots
are much smaller and not so large or blotch-hke. The muscular part of the

Gastrophryne carolinensis 471

tail is colored like the back and has usually along the middle a prominent
horizontal white band which is from 2-7 mm. long. The caudal crests are
more or less translucent with prominent blotches of dorsal color which at
the tip wholly covers the crests. The crests are normally low.

"There are no horny mandibles. On the lower side of the mouth there is
710 labium and in the position of the lower mandible is a fleshy knob not
cornified or differently colored from the surrounding integument. On the
upper side of the mouth there is not a normal labium and in preserved speci-
mens it looks like two more or less independent sideflaps which meet on the
middle line in an emarginate junction.

"The length of the body is contained 1.43-2.8 in the tail, average 1.8.
Width of the body in its own length 1.2-1. 6, average 1.4. Eye lateral, visible
from venter and dorsum, and exactly on the lateral edge formed by the flat-
tened dorsum and venter of the head and forward part of the body. Spiracle
in these spirit specimens undiscoverable and possibly absent in hfe. Anus
median; interorbital space 4.50-7.75 mm., and much greater than the distance
of the eye from the snout (3.25-5.0 mm.). Width of mouth slightly less
than the distance of the eye from snout and 1.6-2.3 in the interorbital dis-
tance. Depth of the tail in its own length 2.2-5.4, the range usually 3.2-4.7,
average 4. Depth of the muscular part at the base of the tail in the depth
of the tail 1.2-1.8 times. Greatest length 37.25 mm. Greatest length of
body 12.75 mm. Greatest length of tail 24.50 mm. Greatest depth of tail 6.0."

In 1920 (p. 32) we wrote "The narrow-mouthed toad, so far as known,
transforms the same season during which the eggs are laid. This period was
formerly considered to be 90 to 100 days, but Deckert's captives required
only 16 days from hatching to transformation, an amazingly short period.
The largest of the tadpoles of this species reach a length of i 2/3 inches and
are very easily distinguished from those of other species. The body is very
flat, and the depth of it is contained i 1/2 times in the width, while other
tadpoles have round bodies; there is no spiracle (a mistake); there are no
horny -edged mandibles, and the lower lip of ordinary tadpoles is not present,
while the upper has either a faint row of teeth or none at all. The color of
the tadpole is quite conspicuous. On the back and sides it is a uniform brown
or olive black. Along the middle of the musculature of the tail there is a
bright, clear, white band one-fourth to one-half inch long. Along either side
of the belly there is a similar white line and most of the belly is of this clear
white. All in all it is our most remarkable tadpole."

In 1923 (p. 34) we have Gastrophryne carolinensis as "black flat tadpoles,
neither mandibles, not labia, nor labial teeth, the spiracle median and just
ahead of anus." By an accident Mrs. Wright discovered the presence of spira-
cle by observation on the development of the hind legs wherein they separate
the two juxtaposed structures, anus and spiracle, the latter going ahead of
the hind legs and the anus remaining behind.

472 Frogs of the Okefinokee Swamp

LARVAL PERIOD

Deckert (19 14, No. 9, pp. i, 2) gives 16 days for the larval period of some
larvae kept in an enamelled pan (see Transformation Size).

The earhest we have taken eggs is May 20, 1921, the earliest transforma-
tion we record is June 25, 1922 or June 12, 19 12. This gives little evidence
but the intervals are 36 days or 23 days. In 192 1 in certain ditches the first
eggs or egg-laying we discovered came May 20. On June 16 we record mature
tadpoles in these ponds 27 days later. On July 23 to 26^ we found our first
tadpoles transformed in these pools. This gives 64 to 67 days from hatching.
In 192 1 the earliest transformation of a few tadpoles came July 7 and many
transformations July 25 and 26. If these were laid May 20-22 this would
give 48 to 50 days from fresh eggs to transformation. In one series of ponds
beside the railroad where eggs were first recorded May 2 2 we first found them
transforming July 23 or 62 days later. We thus have possibilities of 23, 27,
36, 48, 50, 62, 64, 67 days for larval periods, but far removed from Deckert's
control series. He must have had an optimum of conditions to get 20 days
from egg to transformation.

TRANSFORMATION

Period. Deckert's tadpoles transformed September 16 and in 191 2 we
recorded tadpoles we thought would transform in September or October.
If in nature 60 days be allowed for the larval period the eggs we took August
17 would not transform before October 15. In 19 12 we have transformation
June 12 and June 24, July 4. In 192 1 we found transformation from July
7-August I. In 1922 from June 25 to August 25 we took transformed Gas-
trophrijne. For three years it began from June 12- July 7. From mid- June
to mid-October seems the approximate period of transformation.

Size. The first note on transformation comes in Deckert's notes (1914,
No. 9, pp. 1-2): ''Eggs laid, night of August 27, 191 1.

Aug. 28. One egg-mass put in flat enamelled pan in about i 1/2 inches

of water. Larvae straightened out, but still in the egg-mass.
Aug. 30. Larvae released from egg-mass, 1/8 inch long, with small tufts.
Sept. I. Larvae 3/16 inch long, gills disappearing.
Sept. 2. Larvae 1/4 inch long, all have lost gills.
Sept. 8. Hind legs budding, length of larvae 1/2 inch.
Sept. 12. Legs fully developed, arms appearing, tail getting shorter,

larvae 5/8 inch.
Sept. 14. Larvae breathing air, tail is but a short stump,
Sept. 16. Tail gone, the young toads measure 3/8 inch from snout to
vent."

This is development from egg-laying to transformation in 20 days, almost
as rapid as in Scaphiopus holbrookii.

In 191 2 the sizes of "nineteen young transformed narrow-mouthed toads
measure from 11-12 mm., the average is 11. 5 mm., the mode 11. 5 mm."

Gastwphryne carolinensis 473

In 1920 (pp. 34, 36) we stated that "the narrow-mouthed toad tadpole
may transform from the middle of June to September or October. The average
size at transformation is 0.5 of an inch (see Fig. 2 and Plate XXII, Fig. 9)".

In 192 1 we recorded transformation but strangely enough collected but
few. They are: On July 8, 192 1, a series of four transformed frogs amongst
a large series of two-legged tadpoles, measured 12, 12, 12.5 and 13 mm.
respectively. A series of three transformed taken July 23, 192 1, (with
mature tadpoles, tadpoles 20-21 mm. and small tadpoles 12-14 mm.) meas-
ured II, 1 1.5 and 1 1.5 mm. Finally three vials of transformed and tadpole
individuals of July 25, 192 1 (really one lot) were as follows: Vial where they
had dried up 7, 8, 8, 8, 9 mm., another vial with one 9.5 mm. semi-dry, and
another vial not much dry 11 mm. In 1922 we took several series of trans-
formation on June 23, June 24, and July 11. In these forms the tadpoles
with four legs sometimes retained its tadpole mouth until tail stub reached
7-10 mm. in length, or put in the other way the frog mouth sometimes came
in at tail length of 13.5-9 mm., though normally below 10 mm. The meas-
urements of 26 transformed frogs of three series of 1922 are in range from
9-1 1.5 mm., or two at 9 mm., three at 9.5 mm., eight at 10 mm., six at 10.5
mm., six at 11 mm., one at 11.5 mm. This gives a lower range than in 1912,
an average of 10 mm. and mode of 10 mm. The total range then for 191 2-
1922 is 7 or 8-12 mm., average 10.8 mm., mode 11 mm. Some Raleigh N. C,
transformed frogs measure from 11. 5- 12 mm. for six specimens. One might
expect greater variability in size of transformation and doubtless it comes
because of the transient character of many of its breeding places. In one or
two cases from non-Okefinokee localities we have found transformation as
small as 8.5 mm.

General remarks. In 1 9 1 2 we made these notes on the transformed frogs.
They have the dorsal coloration of the adults and on the venter, the under
surface of the hind hmbs, the throat and upper breast are as in the adults
but the belly has more or less of the blotchings and reticulation of the tadpole
belly. In none of the various stages from mature tadpole to transformation
is there any very apparent evidence of a spiracle and the left arm does not
appear before the right. (Later we found the spiracle near the vent).

GROWTH

This species has a size of 8.5-12 mm. at transformation, or an average of
10.8 mm., or mode of 11 mm. We have one record of a specimen taken in
the fall 16 mm. long. Another taken in June 9-10, 19 17, at LaPlace, Ala.,
is 15 mm. On April 25 and 26, 192 1, we took two specimens 17 mm. and 21
mm. cf . We apparently have three groups herein indicated on this small
evidence, a transformation group 8.5-12 mm., an intermediate group 15-17
mm. sexless externally, and a group sexually visible externally. The group
15-17 come from late fall to late spring or early June. Does this represent
8.5-12 mm. specimens June 15-October 15 transformers or 8.5-12 mm. indi-
viduals become 1 5-1 7 mm. by November 1 5-June 1 5 later or a possible maximum
growth of 8.5-5 mm. or minimum of 3 mm. in 5-12 months (winter interven-

474 Frogs of the Okefinokee Swamp

ing). None of our 52 adults examined could be sexed externally if below 21
mm. One therefore seems inclined to extend this 15-17 mm. group to 20 mm.,
and pronounce 21 mm. onward as of the second year group.

On June 11 three females were taken 24, 25 and 28 mm., possibly two
modes or groups. On May 23, 192 1, we took seven adults one group of four
23 mm. cT, 23 mm. 9 , 24 mm. cf , 25 cf , and a group of three 28 mm. cf,
29 mm. 9, 30 mm. 9. On June 23, we took four specimens 24 mm. cf,

27 mm. c^, 28 mm. 9 , 28 mm. cT. On July 4, 192 1, we secured a 24 mm. c?
and 28 mm. 9 . On July 3, 1922, a 23 mm. cf , and a 27 mm. 9 . At least
36 of the 52 adults mentioned are 26-21 mm. Our evidence seems to be
8.5-12 mm. at transformation; 15-20 mm. first year olds; 21-26 or 27 mm.
second year olds; 26 or 27-30 mm. our largest adults.

Examination of some of the U. S. National Museum material (59 frogs)
shows transformed individuals at 11-12 mm. One set of five specimens
(U.S.N.M. 9954, 38781-82 F.B. Meek) from Little Sarasota Bay, Florida,
are 17.5, 18, 18.5, 18.5 and 18.5 mm. respectively. One group from Calcasieu,
La., G. Wiirdemann (U.S.N.M. 3978) gave 17, 17, 17.5, 19, 21, 22, 23 mm.
cf, specimens — two groups (17-19, 21-23). One other group (U.S.N.M.
25420-21) of two specimens, 18.5 mm. and 22 mm. cf represent a sexless
stage and a sexual stage externally. Thus far the above material shows
transformation at 11-12 mm.; a group 17-19 mm.; and a group of 21 or 22
mm. onward, a sexually mature assemblage.

On April 18, 1897, at Wilmington, N. C, Dr. Paul Bartsch collected 5
specimens, (U.S.N.M. 37060-64) three of which were 20 mm. 9 , 20.5 mm.
cf and 20 mm. cf, and two of which were 26 mm. cf and 27 mm. <f. Have
we two model groups? Or on May 23, 19 10, Julius Hurter, Mobile Co.,
Ala., takes two (U.S.N.M. 42548-49) 25 mm. cf and 28 mm. d^. Two days
and five days Later he collects a 28 mm. cT (U.S.N.M. 57698) and another

28 mm. cf (U.S.N.M. 57697). Or May 1 5 at New Orleans, La., they (U.S.N.M.
53188-87) have a 20-5 mm. cf and 27 mm. 9 .

Or on May 25, 1891, J. Hurter takes in Missouri two females (U.S.N.M.
57700-01) 21.5 mm. and 31 mm. respectively. These selected instances show
11-12 mm. at transformation, 17-19 mm. an intermediate group, a 20-25 mni.
group and a 26-31 mm. group. A plotting of the whole 59 frogs reveal 11-12
mm. at transformation, 17-19 mm. apparently first year olds, 20-25 mm.
second year olds, 26-31 mm. three year olds. We have no Okefinokee speci-
means at 20 mm. The U. S. National Museum material shows sex externally
at that size (U.S.N.M. Nos. 37062-64, 53188) and our specimens at 21 mm.
Our largest Okefinokee specimen is 30 mm., a female, but the U. S. National
Museum collection has several females from 32 (U.S.N.M. 57702, 37o7)-36
mm. (U.S.N.M. 57614, 57703-4). We have in our collection four specimens
from Raleigh, N. C, August 1893, namely, 26 mm. cT, 26 mm. 9, 27 mm.
cf and 32 mm. 9 . Is the last of a different mode? Are all these 32-32 mm.
females more than three year olds? The whole evidence seems to be 8.5-12
mm. at transformation, 15-19-20 mm. for first year olds, 20 or 21 mm.-25 or

Gastrophryne carolinensis 475

27 mm. for two year olds, 25 or 27 mm.-3i mm. for three year olds. Possibly
the 32-36 mm. specimens should be three year olds, making that group
27-36 mm.

ENEMIES

On June 5, 191 2, we took one garter snake (Thamnophis sirtalis) which
had eaten this species.

AUTUMNAL DISAPPEARANCE

Our records range from March to September. In the U.S.N.M. collec-
tions we find none taken after September. Surely this species must remain
out until November at least.

AFFINITIES

Holbrook describes this species in 1836 (p. 83). Later Dumeril and Bibron
describes another form E. rugosum (1841, p. 744) which today is accredited
to G. carolinense. Holbrook in his second edition (1842, Vol. V, p, 25) rather
testily writes: "Dumeril and Bibron describe an Engystoma rugosa as inhab-
itating the United States, and suppose that I have confounded it with the
Engystoma carolinense. Now I never saw their Engystoma rugosa, and if I
had, should never have mistaken it for the Engystoma carolinense, which has
a smooth skin. The only ecaudate batrachian animal, with which I am
acquainted, resembling an Engystoma in form and size, is the Bufo quercicus;
but this is a true toad, with parotid glands, warty skin, etc., whereas all the
genus Engystoma, as I received it, have smooth skins and no parotid glands.
Their Engystoma rugosa is probably a Mexican animal, as they say it came
from 'des parties meridionales de FAmerique du Nord.' "

In May, 1854, C. Girard (1856, p. 88) writes: "15 Engystoma rugosum
Dum. and B. erp. gen. VIII, 1841, 744. Said to occur in the same regions
as the preceding species {carolinensis). Have never observed it, and there-
fore cannot endorse it as a North American species."

Hallowell (1856, p. 252) because the Academy specimens and Holbrook's
plate of E. carolinensis appear brown or chestnut above and his lone specimen
from Nebraska and Kansas appears olive, described a new form, Engystoma
olivaceum. Furthermore he held Holbrook gives the distribution of E. caro-
linense as from Charleston to the lower Mississippi valley.

In 1859 (pp. 169, 170) Chas. Girard described Engystoma texense, "This
species is allied to E. carolinense, and differs from it by a more depressed and
flattened head, a more truncated snout, which, as usual, protrudes beyond
the lower jaw. The body itself is Hkewise more depressed, and the limbs
assume a slender appearance.

"The head is continuous with the body, and constitutes about the third
of their combined length. The cleft of the mouth does not extend as far back
as in E. carolinense, since it corresponds to a perpendicular line drawn behind
the pupil. The longitudinal diameter of the eye is equal to the distance
between the orbit and the nostril. The interocular space, measured across

476 Fi'ogs of the Okefinokee Swamp

the interior rim of the orbits, is greater than the rostral space from the orbits
forwards. The symphysis of the lower jaw presents the same structure as
in the species just alluded to.

"The skin is perfectly smooth throughout in all the specimens which we
have examined; they were collected in April, and are, no doubt, hable to as-
sume a rougher appearance during the hotter and more dry season of the year.

"The ground color of the upper region of the head, body and limbs, is of
a light olivaceous brown tint, anteriorly uniform, posteriorly besprinkled
with small black spots or dots especially over the coccyx and thighs. A
whitish tint prevades uniformly throughout the inferior region.

"Specimens of this species were procured in Texas, by Capt. John Pope."

In 1909 (June 25) John K. Strecker described a new form for the U. S. A.,
namely, Engy stoma areolata Strecker. Dr. Stejneger has noted these peculiar
specimens. Mr. Strecker uses the pustulate upper surfaces, coloration and
short hind feet for some of his characters. Victoria, Refugio County, Texas,
the type locality must be an interesting place. The present author was never
in it but a few hours. If G. texense and G. areolata appear together under
the same log one wonders if stray G. carolinense rarities also in this region
would also domicile in the same place. When the first breeding rain came
would they recognize their respective kinds? By any chance has Gastrophryne
areolata any connection with Engy stoma rugosum (Dumeril and Bibron)
which Boulenger reduced to G. carolinense synonymy? Or are they the
roughened "hotter and more dry season" forms Girard intimates for G.
texense .

In 1925 we from eggs, call, general appearance and tadpoles concluded
that G. texense might be different, yet intensive study and careful comparisons
need to be made. We have before us detailed measurements of the type 20
mm. of Gastrophryne areolata and a 28 mm. cf , measurements of 20 mm. cf,
20 mm. 9 and 28 mm. 9 , Gastrophryne texense and 20 mm. cf , 20 mm. 9 ,
28 mm. cf , 28 mm. 9 and 36 mm. 9 of Gastrophryne carolinensis from the
U. S. National Museum collection, and 28 mm. cf and 28 mm. 9 Gastro-
phryne carolinensis from Okefinokee. These actual measurements do not
reveal as many differences in measurements as some of the descriptions might
lead one to believe. The Gastrophryne have too many "relative" characters
not definitely defined, if they can be.

The pustular or rough or tuberculate character is variable. Of one mated
pair captured July 3, 192 1, in the center of the swamp each is very smooth
but a mated pair taken one year later on the esastern edge of the swamp has
the female rough to the ridge between the eyes and the male rough for a short
distance ahead of the vent. A collection of 7 adults taken May 23, 192 1, on
Billy's Island separate on larger size and roughness into females (2) and
smaller size and smoothness into males (5). The one rough or tubercular
character is not a sexual character entirely, nor a wet or dry season character
solely. Gastrophryne needs careful study.

Gastrophryne carolinensis 477

BIBLIOGRAPHY

1 89 1 Boulenger, G. A. Ann. Nat. Hist. ser. 6, v. 8, p. 453.

1896 Brimley, C. S. Batrachia Found at Raleigh, N. C. Am. Nat. XXX, June 1896,
No, 354, p. 501.

1907 . A Key to the Species of Frogs and Toads Liable to Occur in North Caro-
lina. Journ. Elisha Mitchell Sci. Soc, Vol. XXIII, No. 4, Dec. 1907, pp. 157, 159

1026 . Revised Key and List of the Amphibians and Reptiles of North Carolina,

Journ. Elisha Mitchell Sci. Soc, Vol. 42, Nos. i & 2, Oct. 1926, p. 81.

1889 Cope, E. D. The Batrachia of North America, Bull. No. 34, U. S. Nat. Mus. 1889
PP- 385, 386.

19 13 Deckert, R. F. List of Salientia from Near Jacksonville, Florida. Copeia, Feb
14, 1914, No. 3, p. 3.

19 1 5 . Further Notes on the Salientia of Jacksonville, Florida. Copeia, Aug. 29,

1914, No. 9, pp. 1-3.

192 1 . Amphibian Notes from Dade Co., Florida. Copeia, Mar. 15, 192 1, No. 92

1922 . Notes on Dade County Salientia. Copeia, Nov. 1922, No. 122, p. 88.

1842 DeKay, J. E. Zoology of New York, Part III, 1842, p. 65.

1841 Dumeril, A. M. C. et G. Bibron. Erpetologie Generate Vol. 8, pp. 743, 744.

1918 Dunn, E. R. A Prehminary List of the Reptiles and Amphibians of Virginia. Copeia,
No. 53, Jan. 25, 1918, p. 22.

1892 Garman, H. A Synopsis of the Reptiles and Amphibians of Illinois. Bull. 111. State
Lab. Nat. Hist., Vol. Ill, pp. 331-333.

1856 Girard, Charles. A list of the North American Bufonids, with Diagnosis of New
Species. Proc. Acad. Nat. Sci. Phila. Vol. VII 1854. 1855 Phila. 1856, p. 88 May 1854.

i860 Girard, Charles. Herpetological Notices. Proc. Acad. Nat .Sciences, Phila., 1859,
pp. 169, 170.

1858 Gunther, A. Catalogue of the Batrachia Salientia Brit. Museum, 1858, pp. 51,52.

1856 Hallowell, E. Reptiles from Kansas and Nebraska Proc. Acad. Nat. Sci. Phila.,

Oct. 1856, p. 252.

1842 Holbrook, J. E. North American Herpetology, Vol. V, 1842, pp. 23, 25.

1924 Holt, E. G. Additional Records for the Alabama Herpetological Catalogue. Copeia,
No. 135, Nov. 18, 1924, p. 95.

1893 Hurter, Julius. Catalogue of Reptiles and Batrachians Found in the Vicinity of St.
Louis, Mo. Trans. Acad. Sci. St. Louis, Vol. VI, No. II, Dec. 12, 1893, p. 253.

1897 Hurter, Julius. A Contribution to the Herpetology of Missouri. Trans. Acad.
Sci. St. Louis, Dec. 31, 1897, p. 503.

1855 Le Conte, John. Decsriptive Catalogue of the Ranina of the LTnited States. Proc.

Acad. Nat. Sci. Phila., Dec. 1855, pp. 430, 431.
1927 Lindsdale, J. M. Amphibians and Reptiles of Doniphan County, Kansas. Copeia,

No. 164, July-Sept. 1927, p. 77.

1922 Loding, H. P. A Preliminary Catalogue of Alabama Amphibians and Reptiles.
Mus. Paper No. 5, Ala. Mus. Nat. Hist. Geol. Survey Ala., Sept. 1922, p. 21.

1894 Loennberg, Einar. Notes on Reptiles and Batrachians Collected in Florida in 1892
and 1893. Proc. U. S. N. M., Vol. XVII, 1894, 1895, p. 338.

1924 Myers, G. S. Amphibians and Reptiles from Wilmington, N. C. Copeia, No. 131,

June 30, 1924, p. 60.
1926 Nieden, Fr. Das Tierreich Anura II, pp. 64, 65.

1 9 19 Pope, Philip H. Some Notes on the Amphibians of Houston, Texas. Copeia, Dec.
31, 1919, No. 76, p. 94.

1920 Potter, Doreen. Copeia, Sept. 16, 1920, No. 86, p. 82.

1895 Rhoads, Samuel N. Contributions to the Zoology of Tennessee. No. i. Reptiles
and Amphibians. Proc. Acad. Nat. Sci. Phila. 1895, 1896, Part I, Jan. -March, p. 396.

1924 Schmidt, K. P. A List of Amphibians and Reptiles Collected near Charleston, S. C.

Copeia, No. 132, July 15, 1924, p. 68.
1909 Strecker, John K. Notes on the Narrow-mouthed Toads (Engystoma) and the

Description of a New Species from Southeastern Texas. Proc. Biol. Soc. Wash.,

June 25, 1909, Vol. XXII, pp. 1 15-120.
1915 . Reptiles and Amphibians of Texas. Baylor Bulletin Vol. XVIII, No. 4,

Aug. 1915, pp. 46, 47.
1920 Wright, A. H. Frogs: Their Natural History and Utilization. Rept. U. S. Com.

Fisheries, Appendix VI, Doc. 888, pp. 29, 30.

1923 \ — • The Tadpoles of the Frogs of Okefinokee Swamp, Georgia. The Ana-
tomical Record, Jan. 20, 1923, p. 406.

1923 . The Sahentia of the Okefinokee Swamp, Georgia. Copeia, Feb. i, 1923,

No. 115, p. 34-

1926 . The Vertebrate Life of Okefinokee Swamp in Relation to the Atlantic

Coastal Plain, Ecology, Vol. VII, No. i, Jan. 1926, p. 83.

1924 Weight, A. H. and A. A. Wright. A Key to the Salientia East of the Mississippi
River. The American Naturalist, Vol. LVIII, July- August, 1924, pp. 378, 379.

MAP
OKEFINOKEE SWAMP

Plate XL\

INDEX

Abastor erythrogrammus, io8

Abbott, C. C, 74, 8i, 82, 83, 87, 88, 97, 99,
102, 106, 161, 164, 166, 167, 168, 177, 179,
184, 185, 186, 188, 189, 190, 194, 228, 234,

235, 247

Acad, Nat. Sci. Phila., 228, 245, 449, 475
Acadian flycatcher, 314
Acantharchus pomotis, 238, 447
Acer rubrum, 238

Acknowledgments, 1-3

Acris, 13, 26, 29, 30, 50, 65, 68, 117, 191,
192, 193, 194. 202, 203, 211, 213, 220, 225,
272, 282, 289, 304, 305, 330, 369

Acris acheta, 191

.4ms crepitans, 68, 156, 157, 164, 166, 168,

191, 192, 193, 194
Acris gryllus, 3, 4, 12, 13, 16, 18, 19, 24, 25,
26, 29, 30, 31, 32, 33, 34, 35, 37, 38, 39, 40,
50, 57, 58, 59, 60, 61, 63, 64, 66, 67, 68, 92,
124, 146, 156-196, 202, 211, 225, 226, 229,
235, 237, 257, 261, 262, 289, 290, 303, 309,
314, 316, 324, 461

Acris gryllus bufonia, 192, 194

Acris gryllus crepitans, 165, 168, 192, 193

Acris gryllus gryllus, 4

Acris nigritus, 194

Acris pickeringii, 192, 194

Adirondack Mts., N. Y., 397, 400, 402, 403,
404, 405, 407, 408, 409, 410, 413, 414, 418,
438, 449, 451

Affinities, 62-67, S. holbrookii, 105-106; B.
quercicus, 131, 132; B. terrestris, 153, 154,
1555 '4. gryllus, 190-194; P. nigrita, 202,
203; P. ocularis, 225, 226; H. andersonii,
245-247; H. cinerea, 271; H. femoralis,
293, 294; H. gratiosa, 309, 310; H.
squirella, 324; H. versicolor, 333; R.
aesopus, 350-352; R. grylio, 382, 383; R.
heckscheri, 396, 397; R. septentrionalis,
415-418; R. sphenocephala, 434; R.
virgatipes, 452; G. carolinensis, 475, 476

Alabama, 15, 107, 108, 113, 133, 157, 196,
204, 229, 248, 257, 272, 297, 310, 312,
314, 325, 365, 375, 419, 452, 453, 454, 473,
474, 477

Albmo frogs, 353, 392

Alders, 407

Alexandria, Va., 255

Algonquin Park, Can., 400

Allapaha, Ga., 4, 204, 272

Allard, H. A., 140, 155

Alleghany Mts., 193

Allen, A. A., 199

Allen, J. A., 76, 106

Allen, M. J., 385

Alligator Swamp, Fla., 384, 386, 393

Alnus incanus, 407

Altamaha River, Ga., 272

Ambystoma, 286, 466

Ambystoma angulatum, 10

Amby stoma conspersum, 6

Amblystoma opacum, 6

Amblystoma punctatum, 6

Amblystoma talpoideum, 6

Am. Mus. Nat. Hist., 3, 228, 318, 323, 348,
349, 384, 398

American spade-foot, 106

Ames, Iowa, 157, 174, 179

Amphiuma means, 4, 429

Anderson, R. D., 448

Anderson, S. C, 228, 240, 435, 453

Anderson tree frog, 28, 31, 235

Anderson tree toad, 227

Anderson's Hyla, 227, 234

Appel, W. D., 449

Arch Creek, Fla., 314

Aristook Co., Me., 409

Arizona, 33

Arkansas, 42, 73, 74, 206, 325, 352, 454

ArUngton, Fla., 207, 348, 350

Armstrong, A. J., i, 9

Aronia, 116

Arrowhead, 386

Arrow arums, 371

Ascaphus truei, 66

Aschemeier, C. R., 209, 213, 217

Athabaska, 201

Atkinson, C., 162, 194

Atlanta, Ga., 453

Atlantic City, N. J., 435

Atlantic Co., N. J., 235

Atlantic coastal plain 107, 108, 133, 196,
419, 452, 457

Atsion Creek, N. J., 235

Auburn, N. Y., 397

Auburndale, Fla., 114, 337, 348

Austroriparian region, 134, 155, 228, 351

Autaugaville, Ala., 114

Autumnal disappearance, 24; S. 'holbrookii,
104-105; B. quercicus, 131; B. terrestris,
153; -4- gryllus, 188; P. nigrita, 202; P.
ocularis, 225; H. andersonii, 245; H.
cinerea, 270, 271, 293; H. femoralis, 293;
H. gratiosa, 309; H. squirella, 323, 324;
H. versicolor, 333; R. aesopus, 350; R.
clamitans, 364; R. grylio, 382; R. spheno-
cephala, 434; R. virgatipes, 451; G. caro-
linensis, 475

Axillary amplexus, 28, 29, 30

Azalea viscosa, 238

B

Babcock, H. L., 157
Bailey, V., 114, 206
Bainbridge, Ga., 9

Baird, S. F. IX, 5, 6, 106, 168, 191, 192, 198,
203, 204, 205, 207, 227, 234, 247, 351, 408
Baker County, Fla., 10, 392
Baldwin Co., Ala., 452
Baltimore, Md., 163, 245

479

48o

Frogs of the Okefinokee Swamp

Baltrusol, N. J., 157

Bamboo, 212

Banded Hyla, 248

Barachias, Ala., 310, 453

Barbec 322

Barber, C. M., 170

Barbour, F. K., 236, 244, 450

Barbour, T., 3, 33, 62, 73, 107, 133, 196, 203,
227, 236, 239, 243, 244, 245, 247, 248, 255,
271, 272, 294, 325, 334. 348, 349, 352, 364,
365, 368, 370, 381, 383, 384, 397, 419,
423, 434. 435, 437, 450, 452

Barking frog, 294, 297, 300, 301

Barrens, 12

Barton, B. S., 5

Bartram, Wm., 133, 138, 155, 161, 191,
372, 384, 422

Bartram's Botanic Garden, 333

Bartsch, Paiul, 474

Bat-fish, 340

Batrachus tau, 468

Batsta Creek, N. J., 235

Battle, P., 9

Bawling frog, 352

Bay St. Louis, Miss., 114, 134, 157, 189, 196,
199, 206, 253, 254, 312, 364, 365

Bay swamp, 10

Bays, 12, 224

Bean, B. A., 398, 413

Bean, T. H., 309, 350

Beaufort, N. C, 113, 450

Beaumont, Tex., 248

Beaver, 400

Beecher Pt., Fla., 252

Beeville, Tex., 157, 205, 227, 248, 325, 326

Bejaria racemosa, 438

Bellfrog, 14, 248, 256

Bellair, Fla., 134

Belly bumper, 352

Berwick, La., 248

Bibliography, S. holhrookii, 106-107; B. quer-
cicus, 132, 133; B. terrestris 155, 156; A.
gryllus 194-196; P. nigrita 204; P. ocularis,
226-227; H. andersonii, 247-248; H.
cinerea, 271; H. femoralis, 294; H. grati-
osa, 310; H. squirella, 324; H. versi-
color, 333; R. aesopus, 352; R. clamitans,
364; R. grylio, 384; R. Iieckscheri, 397;
R. septentrionalis, 418-419; R. spheno-
cephala, 434, 435; R. virgatipes, 452; G.
carolinensis, 476, 477

Bidens, 163

Biloxi, Miss., 157, 312, 385, 458

Bishop, S. C, I, 9, II, 130, 132, 397, 402,
403, 405, 406, 407, 408, 409, 410, 414

Black chorus frog, 13

Black cricket frog, 57

Black frog, 352

Black-nosed dace, 407

Blacksburg, S. C, 453

Black snake, 152, 190, 270, 292, 309, 434

Black spotted tree frog, 196

Black swamp cricket frog, 28

Black toads, 138

Bladderwort, 264, 368, 443

Blatchley, W. S., 163, 167, 194

Bob white, 437

Boleichthys fusiformis, 447

Bonnaterre, J. P., IX, 13, 133, 134

Bonnets, 163

Boone's Pond, N. C, 170

Borachio, Texas, 156

Bosc, L. A. G., 5, 154, 225, 312, 456

Boston, Mass., 87

Boulenger, G. A., V, 29, 40, 41, 65, 82, 95,

106, 120, 131, 135, 155, 165, 192, 194,
200, 203, 204, 207, 208, 226, 245, 293,

348, 349, 364, 365, 371, 383, 384, 397.

416, 418, 420, 434, 442, 449, 451, 452,

454, 455. 458, 476, 477
Boyle, H. S., 162, 194
Bradley, J. C, 9, 157, 185, 196, 204, 262,

272, 276, 292, 322, 365, 370, 423, 432
Brady, M. K., 33, 188
Branch swamps, 1 1
Brantley County, Ga., 10
Brasenia, 368, 400
Breeding sizes, 18, 24-26, 58, 59, 63
Brentwood, Fla., 337
Brevard, N. C, 169
Brevicipitidae, 28, 41, 57
Brickells Hammock, Fla., 196
Brimley, C. S., 4, 75, 83, 88, 92, 95, 106,

107, 108, 112, 132, 133, 134, 135, 144,
155, 157, 177, 188, 193, 194, 196, 199,
206, 207, 208, 226, 227, 228, 229, 247,
248, 253, 254, 271, 272, 294, 295, 311,
324, 419, 434, 435, 449, 450, 452, 454,
464, 465, 477

Brimley, H. H., 253, 254, 449

Brook trout, 415

Brown snake, 252

Brownsville, Texas, 226

Brownton, N. J., 229

Bruce Co., Ont., 397

Bufo, 13, 26, 29, 64, 65, 85,''88, 98, 99, 106,

108, 109, 127, 132,4141,4145, 150, 270,
424, 461, 466, 470

Bufo americanus, 5, 29, 33, 34, 36, 43, 59,
64, 66, 68, 71, 114, 120, 125, 131, 134,
135, 139, 140, 147, 150, 153, 154. 155,
353-

Bujo calamita, 65, 452

Bufo cognatus, 65, 115, 116, 132, 155

Bufo compactilis, 64, 65, 66, 115, 116, 127,
132

Bufo debilis, 129, 132

Bufo dialophus, 131

Bufo erithronatus, 131

Bufo erythronatus, 5, 138, 139, 155

Bufo fowleri, 15, 16, 29, 34, 36, 43, 44, 63,
64, 65, 66, 68, 71, 109, 134, 140, 144,
153, 154, 155, 176, 237, 272, 349, 452

Bufo frontosus, 155

Bufo gibbosus, 456

Bufo halophila, 206

Bufo hemiophrys, 155

Bufo lentiginosus, 3, 4, 131, 132, 137, 139,
140, 148, 153, 154, 155, 349, 456

Bufo lentiginosus mnericanu^, 6, 131, 147

Bufo lentiginosus lentiginosus, 4, 134

Bufo lentiginosus musicu^, 135

Bufo lentiginosus woodhousei, 132

Bufo musicus, 138, 139, 155

Bufo punctatus, 42, 132

Bufo quercicus, 3, 4, 12, 13, 15, 18, 19, 24,
25, 26, 29, 31, 32, 33, 34, 36, 37, 38, 39,
57, 58, 59, 60, 61, 62, 63, 64, 65, 66, 68,

Index

481

107-133, 148, 155, 199, 215, 216, 261,
265, 278, 282, 294, 303, 304, 316, 319,
336, 349, 350, 461, 467,475

Bufo terrestris, 12, 13, 16, 17, 18, 19, 21,
22, 23, 25, 26, 29, 30, 32, 33, 34, 36, 37>
38, 39, 40, 42, 57, 58, 59, 60, 61, 62, 63,
64, 65, 66, 68, 71, 109, no, in, 113, 119,
123, 125, 128, 176, 294, 299, 316, 336,
460, 461

Bufo valliceps, 65, 66, 68, 252

Bufo vulgaris, 154

Biifo woodhousei. 154, 155, 349

Bufonidae, 28, 33, 42, 57, 477

BuUfrog, 7, 8, 17, 28, 38, 39, 176, 213, 248,
352, 364, 370, 382, 397, 399, 403, 407

Bully, 352

Burlington Co., N. J., 235, 439, 443

Burmannia, 443

Butler Co., Mo., 454, 457

Buttonbush, 163, 370, 392

Cahn, A. R., 9

Calamita arbor ea, 271

Calcasieu Parish, La., 454, 474

Calhoun Co., Ala., 452

Cahco bush, 277

California, 205

Callahan, Fla., 94, 129, 172, 259, 272, 291,

316, 384, 386, 393, 394
Caloosatcha River, Fla., 157
Cambridge, Mass., 76, 77, 95, 107
Camden Co., Ga., 10
Camden, Co., N. J., 234
Camp, C. L., 228
Camp, R. D., 252
Canada, 49, 50, 155, 166, 197, 201, 325,

352, 397, 400, 451
Canadian Northwest, 156
Cape May, N. J., 89
Cape May Co., N. J., 439
Cape Sable, Fla., 310, 419
Carnegie Museum, 3
Carolina, 73, 133, 134, 152, I54, 190, 191,

196, 197, 228, 248, 251, 272, 275, 310,

372, 456
Carohna chickadee, 229, 314
Carolina Co., Va., 453
Carolina Hyla, 248
Carolina toad, 133

Carohna tree frog, 62, 248, 268, 293, 452
Carohna wren, 314
Carpenter frog, 14, 28, 31, 38, 58, 63, 74,

403, 435-452
Carteret Co., N. C, 107
Cary, N. C, 330, 453
Cassandra, 400
Castalia, 262
Caswell, Me., 399
Catalogue of species, 13-15
Catbriar, 371

Catesby, Mark, 138, 152, 154
Catfish, 394

Catskill Mts., N. Y., 397, 398, 451
Cedar Cr., Fla., 337
Cedar Grove, N. J., 439
Cedar Swamp Cr., N. J.. 439
Centrarchidae, 212
Cephalanthus occidentalis, 370, 392

Ceralophris, 105

Ceratophrys, 135

Chamaecyparis thuyoides 238

Chameleon Hyla, 325

Champaign Co., 111., 188

Chapel Point, Md., 157, 185

Chandler, A. C, 157

Chaney, L. W., 81

Chapin, J., 228, 229, 240, 243, 244, 435

Charleston, S. C, 107, 133, 138, 207, 248,

310, 323, 324- 452, 475, 477
Charlotte, N. C, 229, 453
Charlton Co., Ga., 10
Charming toad. 133, 134
Chattanooga, Tenn., 457
Check list (Stejneger & Barbour) .S'. hol-
brookii, 73; B. quercicus, 107; B. terrestris,
133; A. gryllus, 156; P. nigrita, 196; P.
ocularis, 206; H. andersonii, 228; H.
cinerea, 248; H. femoralis, 272; H.
gratiosa, 294; H. squirella, 310; H. versi-
color, 325; R. aesopus, 334; R. clamitans,
352; R. grylio, 364; R. septenlrionalis, 397;
R. sphenocephnla, 319; R. virgatipes, 435;
G. carolinensis, 452
Chelopus guttatus, 437
Chelydra serpentina, 245
Cheraw, S. C, 158, 228, 240, 259, 272, 435
Cherokee Co., Ala., 452
Chesser, B., i, 441
Chesser, H., i, 276, 451
Chesser, R., i, 74, 445
Chesser, R. A., i, 283, 299, 300, 441
Chesser, S., i
Chesser, T., i

Chessers, 2, 283, 299, 303, 440, 445
Chicken turtle, 309, 323
Chickopee, Mass., 77
Chilophryne dialopha, 131
Chmophilus, 7, 117, 171, 176, 183, 203, 205,

212, 215, 216, 220, 225, 277, 374
Chorophilus angulatus, 207
Chorophilus coiii, 204, 210
Chorophilus feriarum, 198, 203
Chorophilus nigritus, 3, 5, 6, 196, 197, 203,

225
Chorophilus nigritus mgritus, 198
Chorophilus occidentalis, 3, 4, 6, 196, 204,

205, 206, 210
Chorophilus ocularis, 3, 205, 215, 216, 299
Chorophilus ornatus, 4, 6, 204, 205, 219,

225
Chorophilus triseriatus, 182, 198, 203
Chorophilus verrucosus, 203
Chorus frog, 57
Chrosomus erythrogaster, 407
Chrysemys picta, 66, 245, 437
Chub, 415

Cimarron Co., Okla., 74
Cinereous frog, 248
Claa?sen, P. W., 397, 402
Clarcona, Fla., 454, 457
Clark, H. W., 413
Clarkstown, N. Y., 73

Clear Lake, N. Y., 397, 402, 403, 410, 414
Cleared and cultivated fields.iii, 113, 122,

312
Clematis, 407
Clementon, N. J., 232

482

Frogs of the Okefinokee Swamp

Clemmys guttata, 66, 245

Clemmys muhlenbergii, 66

Clethra alnifolia 183, 238, 370, 392, 443

Cliff Cave, Mo., 454, 457

Cleveland, Tex., 454

Clinch, Co., Ga.. 10

Clyde River, Vt., 398

Cnemidolphorus sexlineatus, 350

Coat bets, 129, 294, 299, 300, 303

Cochran, Miss D., V

Cocoanut Grove, Fla., 314

Colorado, 85, 157, 189, 195

Colorado College, 85

Colorado Springs, Colo., 85

Coloration from life, 27

Coloration of spirit specimens (1912). 16,
B. quercicus, 109; B. terrestris, 135; A.
gryllus, 158, 159; P. nigrita, 197-198; P.
ocularis, 208-210; H. andersonii, 230;
H. cinerea, 249, 250; H. femoralis, 273;
H. gratiosa, 295; R. clamitans, 353; R.
grylio, 365, 366; R. heckscheri (1922),
385; R. sphenocephala 420, 421; G. caro-
Unensis, 454, 455-

Coluber constrictor, 292, 434

Columbia, S. C, 157, 248, 272

Columbus, Ga., 454

Common leopard frog, 420

Common names, *S. holbrookii, 73; B. quer-
cicus, 107; B. terrestris, 133; A. gryllus,
156; P. nigrita, 196; P. occidentalis, 204;
P. ocularis, 206; P. ornata, 227; H.
andersonii, 227; H. cinerea, 248; H.
femoralis, 272; H. gratiosa, 294; H.
squirella, 310; H. versicolor, 325; R. ae-
sopus, 334; R. clamitans, 352; R. grylio,
364; R. heckscheri, 384; R. septentrionalis,
397, R. sphenocephala, 419; R. virgatipes,
435! G- carolinesis, 452

Common shiner, 407

Common spotted frog, 14

Common toad, 88, 103

Common tree frog, 325

Common tree toad, 28,31, 184,295,325-333

Conn, H. W., 81

Connecticut, 78, 107

Connerv Pond, N. Y., 397

Cook, G. H., 234

Cope, E. D., IX, 4, 6, 14, 82, 93, 106, 108,
114, 131. 132, 134, i35r 153, i54r 155.
156, 162, 164, 165, 167, 168, 192, 193,
194, 196, 198, 200, 203, 204, 206, 208,
209, 213, 226, 228, 229, 245, 247, 272,
274, 294, 295, 310, 324, 327, 333, 334,
343, 350, 351, 352, 383, 397. 414, 416,
418, 419, 434, 435, 436, 437, 438, 449.
450, 451, 452, 455, 458, 477

Cope's frog, 194, 403, 435, 444, 445

Coram, N. Y., 86

Corn. Univ. Mus., 380

Couper, J. H., 4, 5

Cow bell frog, 31, 248

Cranes, 190

Craven Co., N. C, 107, 207, 229, 272, 449

Crawfish, 190, 336

Crest of ovulation, 32

Cricket frog, 13, 28, 92, 124, 156-196, 200

Cricket toad, 156

Crosby, C. R., 9, 62, 68, 130, 132, 397, 402,

405

Cross embraces, 28, 29

Cross Lake, Me., 409

Cultivated fields, 11, 262, 299

Cumberland Co., N. C, 134

Cumberland plateau, 193

Cuming, Mr., 197, 227

Cuthbert, Ga., 157

Cut lips, 407

Cuvier, G. L. C. F. D., 75

Cypress bay, 11, 12, 14, 172, 207, 212, 215,
224, 252, 262, 276, 282, 283, 352, 356,
365, 375, 422, 424, 426, 427, 461

Cypress heads, 11, 172, 272

Cypress ponds, 11, 12, 13, 14, 92, 117, 140,
157, 163, 171, 172, 190, 212, 215, 216,
223, 224, 252, 262, 277, 283, 303, 340,
356, 365, 369, 375, 380, 422, 424, 426,
427, 464

Cystignathidae, 334

Cystignathus, 203

Cystignathus nigritus, 5, 191, 197, 198, 200,
204

Cystignathus ornatus, 5, 200, 204, 227

D

Dade Co., Fla., 89, 106, 207, 310, 312, 313,
323, 368, 370, 419, 427. 452, 457, 477

Dahlgren, U., 88

Dane Co., Wise, 162

Danvers, Mass., 86

Dare Co., N. C, 248

Daudin, F. M., 154, 190, 225, 246, 253,
256, 271, 313,456

Dauphin, Ala., 312

Davis, N. S., Jr., 351

Davis, W., I

Davis, W. T., 84, 106, 228, 231, 233, 236,
239, 240, 243, 244, 245, 247, 248, 435,
436, 437, 438, 449, 450, 452

Dayton, Tex., 157

Decatur Co., Ga., 157, 262

Deciduous holly, 407

Deckert, R. F., 3, 74, 89, 92, 93, 96, 106,
113, 115, 120, 126, 128, 129, 133, 135,
137, 139, 140, 155, 156, 162, 169, 193,
194, 196, 199, 200, 201, 204, 205, 206,
208, 209, 211, 212, 213, 214, 226, 249,
251, 256, 262, 271, 272, 275, 276, 277,
284, 294, 297, 298, 304, 310, 312, 313,
323, 324, 334, 338, 341, 342, 347, 349,
352, 365, 368, 370, 382, 384, 419, 423.
427, 428, 434, 452, 457, 459, 460, 464,
466,469,472,477

Decodon, 262

Deirochelys reticularia, 293, 309, 323

De Kay, J. E., 73, 75, 78, 105, 108, 133,
155, 156, 158, 161, 168, 191, 194, 197,
204, 207, 227, 249, 294, 324, 327, 333,

453, 477
De Kay's snake, 252
Del Rio, Tex., 156
Dennisville, N. J., 439
Departiu-es from typical amplexus, 29-30
Dernehl, P. H., 162, 165, 195
Desmognathus fusca, 4, 6
Desmognathus fusca auriculata, 4, 6
Desmognathus nigra, 6

Index

483

De Terville, 456

Devers, Tex., 248, 263

Diadophis punctatus, 379

Dickerson, Miss M. C., 108, 109, 114, 115,
120, 132, 133, 134, 135, 153, 156, 165,
120, 132, 133, 134, 135, 153, 156, 165,
169, 177, 178, 183, 195, 199, 201, 203,
208, 225, 226, 228, 233, 249, 252, 254,
255, 271, 284, 294, 295, 297, 310, 311,
313, 327, 334, 338, 349, 352, 355, 365,
368, 370, 382, 384, 397, 418, 420, 434,
450, 454, 45.5.

Diemyctylus viridescens, 4

Dinsmore, Fla., 172, 337

Dinwiddle, Va., 157, 176, 177, 179. 325, 330

Dismal Swamp, Va., 229

District of Columbia, 106, 195, 200, 248,
253

Ditmars, R. T., 88, 96, 102, 106, 165, 167,
168, 177, 195

Doniphan Co., Kan., 163, 453, 477

Dorset, Ont., 325, 402, 405, 408, 409, 412,
414

Doucette, Tex., 157

Douglaston, L. I., 105

Drake, C. J., 433

Dreen, 12

Drosera, 443

Dry pine barrens, 11, 163, 336, 352

Duck grass, 368

Dumeril, A. M. C. and Bibron, G., 90, 105,
106, 191, 194, 195, 197, 202, 203, 204,
293, 324, 458, 475, 476, 477

Dunn, E. R., 89, 169, 189, 193, 195, 255,
271, 435, 452, 453, 477

DupUn Co., N. C, 107

Durham, N. C, 453

Duval Co., Fla., 133, 352

Dwarf toad, 107, 108

Dyke Marsh, Va., 158

E

East Florida Flatwoods, i o

East Gulf Coastal Plain, 108

Eastern States, 73

Eastport, Fla., 384

Edgecombe Co., N. C, 107

Edwards, Sam, i, 2, 9

Egg bands, 31, 36, 37

Egg complements, 33, 35-37

Egg cyUnders, 31, 36, 37

Egg envelopes, 33, 34, 35-37, 64

Egg files, 30, 36, 37, 126

Egg key, 35-37

Egg plinths, 30, 36, 64

Egg size, 25

Egg spheres, 36, 64

Eggs, 33-37, 64

Attachment, S. holbrookii, 95-97; B.

quercicus, 126; B. terrestris, 147; P.

ocularis, 219, 220; H. cinerea, 263, 264;

H. femoralis, 285, 286; H. gratiosa, 304,

305; R. aesopus, 343, 344; R. clamitans,

360; R. grylio, 375; R. sphenocephala, 428;

R. virgatipes, 444; G. carolinensis 465-467

Dangers, 97; B. quercicus, 127, 128;

B. terrestris, 148; P. ocularis, 219, 220;

H. cinerea, 265; H. femoralis, 287; H.

gratiosa, 305; H. squirella, 320; R.

aesopus, 344; R. clamitans, 361; R. grylio,
376; R. sphenocephala, 429, 430; R. vir-
gatipes, 445; G. carolinensis, 467
Egg description; S. holbrookii, 95-97;
B. quercicus, 127; B. terrestris, 147, 148;

A. gryllus, 179, 180; P. ocularis, 220; H.
cinerea, 264; H. Jemoralis, 286, 287; H.
gratiosa, 305; H. squirella, 319, 320; R.
aesopus, 344; R. clamitans, 360, 361; R.
grylio, 376; R. sphenocephala, 428, 429;
R. virgatipes, 444, 445, 446; G. carolin-
ensis, 465-467

Egg mass, 30, 31, 33-37, 64; S. hol-
brookii, 95-97; B. quercicus, 126; B.
terrestris, 147, 148; A. gryllus, 179, 180;
P. ocularis, 220; H. cinerea, 263, 264; H.
femoralis, 285, 286; H. gratiosa, 304, 305;
H. squirella, 319; R. aesopus, 343, 344;
R. clamitans, 361; R. grtjlio, 375, 376;
R. sphenocephala, 428; R. virgatipes, 444;
G. carolinensis, 465-467
Special; P. nigrita, 202; H. andersonii,
240, 241; H. versicolor, 330; R. heckscheri,
390; R. septentrionalis, 409, 410

Elaphe obsoleta confinis, 293

Elassoma, 190, 379

Eldridge, J. S., 89

Eleutheradoctylus ricordii, 15, 18, 19, 66

Ellis, M. M., 157, 158, 166, 174, 189, 195

Emlie, P. H., 238, 243, 441, 447, 448

El Paso Co., Tex., 156

Emporia, Kan., 157

Enemies, 62, 292, 293; B. quercicus, 131;

B. terrestris, 152, 153; A. gryllus, 189-190;
P. nigrita, 202; P. ocularis, 224.; H.
andersonii, 245; H. cinerea, 270; H.
femoralis, 292, 293; H. gratiosa, 309; H.
squirella, 323; R. aesopus, 349, 350; R.
clamitans^ 364; R. grylio, 382; R. heck-
scheri, 396; R. septentrionalis, 415; R.
sphenocephala, 434; R. virgatipes, 450;
G. carolinensis, 475

Engelhardt, G. P., 84

Engystoma, 374, 453, 467, 468, 475

Engystotrm areolata, 453, 476

Engystoma carolinense, 3, 453, 456, 457, 475

Engystoma, elegans, 454

Engystoma microps, 454

Engystoma olivaceum, 475

Engystoma ovale, 454

Engystoma rugosum, 475, 476

Engystoma texense, 453, 454, 475

Engystoma ustum, 454

Enneacanthus, 245, 437, 447

Equisetum, 308

Erimyzon sucetta oblongus, 437

Eriocaulon, 212, 438, 443, 444

Eriocaulon articulatum, 400

Essex Co., Mass., 107

Etowah Co., Ala., 452

Eumeces brevilineatus, 252

Eureka, Fla., 348

Eustis, Me., 397, 399, 413

Everetts Pond, N. C, 228, 237, 259, 330,

435, 437
Evermann, B. W., 188, 213, 398, 413
Exoglossum maxillingua, 407
Explorations to 1914, 3-9
Explorations to 1922, 9-10

484

Frogs of the Okefinokee Swamp

Fair Haven, Conn., 78, 79

Fall croaking, 27

Fall fish, 407

Fall mating, 27

Fall ovulation, 27; Rana sphehocephala, 27

Farmingdale, N. J., 229, 235

Fayetteville, N. C, 133

Femoral Hyla, 272

Fern bogs, 1 1

Fern prairies, 1 1

Fertilization, 26, 27

First Appearance, 24, S. holbrookii, 76, 77;
B. quercicus, 113, 114; B. terrestris, 138;
A. grylliis, 163; P. nigrita, 199; P. ocu-
laris, 212-213; H. andersonii, 232; H.
cinerea, 252; H. femoralis, 276; H.grati-
osa, 297; H. squirella, 312; H. versicolor,
326; R. aesopus, 337; R. clamitans, 355;
R. grylio, 368; R. septentrionalis, 400; R.
sphenocephala, 423; R. virgatipes, 438; G.
caroUnensis, 458

First Connecticut Lake, N. H., 398, 413

Fisher, G. C, 104, 105, 106

Fitzinger, L. J., 191, 203

Flat Pine Barrens, 10

Flatwood, Ala., 157, 248, 257

Fletcher Lake, Ont., 400

Floating heart, 368

Florida, 10, 15, 16, 41, 42, 44, 46, 47, 49,
50, 52, 53, 54, 55, 56, 66, 73, 74, 93, 94,
107, 108, no, 113, 125, 133, 137, 138,
139, 155, 156, 157, 158, 162, 165, 169,
190, 191, 192, 194, 195, 196, 199, 200,
201, 203, 204, 205, 206, 207, 209, 211,
212, 213, 217, 223, 226, 229, 248, 252,
254, 259, 262, 267, 271, 272, 275, 294,
295, 309, 310 312, 314, 315, 316, 318.
322, 325, 334, 337, 340, 342, 343, 347,
350, 351, 352, 364, 365, 368 370, 372,
375, 382, 383, 384, 385, 386, 387, 393,
419, 423, 427, 428, 451, 452, 453, 454,

474, 477

Florida blue herons, 379

Florida district, 334, 351

Florida forms, 10, 15, 66, 113, 351

Florida frog, 334

Florida gopher frog, 334, 352

Florida grackles, 190, 379, 396, 435

Florida Hyla, 294

Florida Parishes, La., 108, 275, 294

Florida Keys, 15

Floridan subregion, 134, 334

Florida tree frog, 14, 28, 30, 31, 92, 93,
124, 294

Florida tree toad, 297

Food, 62, S. holbrookii, 103-104; B. quer-
cicus, 130; B. terrestris, 152; A. gryllus,
189; P. ocularis, 224; //. andersonii, 244;
H. cinerea, 268-270; H. fenwralis, 292;
H. gratiosa, 309; H. squirella, 323; R.
aesopus, 349; R. clamitans, 364; R. grylio,
382; R. septentrionalis, 415; R. spheno-
cephala, 433; R. virgatipes, 450

Force, Miss E. R., 325, 333, 419, 433, 434

Fort George Island, Fla., 133

Fort Meade, Fla., 134

Fort Ripley, Minn., 397

Foster, Mr. 154

Fountain, P., 7, 8, 9

Fowler, H. W., 84, 88, 89, 106, 162, 167,
169, 195, 196, 199, 204, 228, 234, 248,
327, 333, 352, 437, 438, 442, 450, 452

Fowler's toad, 98, 140, 176

Fox, H., 89

Fox, W. H., 79

Framingham, Mass., 74

Franklin Co., Me., 397, 399, 413

Freneau, N. J., 229

Fried bacon frog, 248, 256

Frog-toad, 452, 464

Frost, S. W., 62, 433

Fruitland Park, Fla., 348

Fry, N. P., 348

Fuertes, L. A., 228, 244

Fulford, Fla., 314

Fulton Lake Chain, X. Y., 397, 400, 409

Fundulus, 153

Funkhouser, W. D., 1,9

Gaige, F. M., 3

Gaige, Mrs. H. T., 162, 195, 336, 352

Gainesville, Fla., 15, 74, 93, 196, 199, 200,

213 ,217, 227, 297, 304, 337, 343, 384, 385.
Gainesville, Ga., 157, 185
Gallberry, 215
Gamhusia, 153, 359, 470
Garman, H., 134, 140, 156, 158, 162, 165,

168, 173, 188, 189, 193, 195, 253, 256,

271, 454, 456, 458, 459, 463, 477
Garnier, J. H., 399, 400, 401, 402, 405, 407,

408, 409, 412, 413, 414, 415, 419
Garrison's Creek, N. Y., 397
Garter snake, 131, 152, 292, 475
Gastrophryne, 13, 26, 29, 65, 92, 278, 299,

360, 453, 460, 461, 472
Gastrophryne areolata, 454, 476
Gastrophryne carolihensis, 12, IT,, 15, 16. 18,

19, 24, 25, 26, 31, 32, 33, 34, 35, 37, 38,

39, 40, 41, 57, 58, 59, 60, 61, 65, 66, 67,

71, 92, 265, 278, 312, 452-477
Gastrophryne texense, 92, 454, 455, 458, 461,

465, 476
Gaylussacia, 116
Gaylussacia dumosa, 199
Gaylussacia frondosa, 277
Geckos, 66
General appearance, S. holbrookii, 75; B.

quercicus, 108, 109; B. terrestris, 134,

135; ^- gryllus, 158; P. nigrita, 196-197;

P. ocularis, 207-208; H. andersonii, 229-;

H. cinerea, 249; H. femoralis, 273; H.

gratiosa, 295; H. squirella, 311; R.

aesopus, 334; R. clamitans, 352; R. grylio,

365; R. hecksheri, 385; R. septentrionalis,

398; R. sphenocephala, 420; R. virgatipes,

435; G. carolinense, 454
General discussion, 15-71
General habits, S. holbrookii, 77-86; B.

quercicus, 114-115; B. terrestris, 139; .4.

gryllus, 164-168; P. nigrita, 199, 200; P.

ocularis, 213; H. andersonii, 233; H.

cinerea, 252-256; H. femoralis, 276-277;

H. gratiosa, 297, 298; H. squirella, 312-

314; H. versicolor, 327; R. aesopus, 337-

339; R. clamitans, 355; R. gnjlio, 368-370;

R. heckscheri, 387; R. sphenocephala, 42;^;

G. caroUnensis, 458, 459

Index

48s

Georgia, 15, 16, 47, 50, 61, 73, 74, 78, 93,
106, 108, 112, 117, 133, 134, 140, 155,
156, 157, 158, 185, 195, 196, 197, 199-
202, 204, 205, 206, 208, 227, 229, 248,
251, 252, 259, 262, 272, 275, 276, 292,
294, 297, 315, 316, 319, 320, 321, 322,
326, 329, 330, 334, 344, 347, 350, 351,
365, 375, 384, 390, 392, 393. 435. 440,
446, 450, 451, 454, 456, 457, 460, 477

Georgia tree frog, 294

Geyer, H., 318, 324

•Gribbes, L., 5

Gibson, L. S., 453

Gill, T. N., V, 29

Gilmore, R. J., 85

Girard, C, IX, 133, 139, 155, 156, 204,
205, 207, 351, 454, 475, 476, 477

Glackens, L. M., 235

Glass snake, 350

Glen St. Mary, Fla., 93, 343, 347

Gloucester, N. C, 157

Godwin, J., i

Godwin, N., i

Golden club, 438

Goose Creek, S. C., 272

Gopher frog, 14, 17, 28, 30, 39, 58, 74, 93,
131, 334-352, 392

Gopher tiu-tle, 14, 336, 337, 349, 352

Gophers, 336

Gopherus polij-phemus, 336

Grasshopper frog, 8

Grassy fields, 1 1

Great Alligator Dismal Swamp, N. C, 229

Great Bear Lake, Can., 197

Great blue herons, 190, 415, 434

Great Egg Harbor River, N. J., 450

Green, J., 5

Green and j'ellow tree-toad, 227

Greenback, 384

Green briars, 231

Green Cove Springs, Fla., 4, 113, 206, 227

Green frog, 14, 17, 28, 31, 38, 58, 352-364,
387, 393, 397, 403, 407, 447, 47o

Green heron, 309

Green tree frog, 14, 28, 248-271, 295, 393

Green tree toad, 248

Greenville, S. C, 196, 197

Greenway, Arkansas, 74, 454

Grey toad, 133

Gronberger, 254, 271

Ground Uzard, 252

Growth, 59-61, S. holbrookii, 103; B. quer-
cicus, 129, 130; B. terrestris, 151, 152;
A. gryllus, 188; P. nigriia, 202; P. ocu-
laris, 224; H. andersonii, 244; H. cinerea,
267, 268; H. gratiosa, 309; H. squirella,
322, 323; H. versicolor, 332; R. aesopus,
348, 349; R. clamitans, 363, 364; R.
grylio, 381; R. heckscheri, 395, 396; R.
septentrionalis, 414, 415; R. spheno-
cephala, 433; R. virgatipes, 449, 450; G.
carolinensis, 473-475

Gulf Coastal Plain, 156, 457

Gulf port, Fla., 348

Gulf States, 157, 325, 365, 379, 382, 457

Gulf Strip, Lower Austral Zone, 10, 272

Gunther, A., 106, 155, 192, 194, 195, 197,
200, 203, 204, 227, 458, 477

Gwinett Co., Ga., 140
Gyrinophilus porphyriticus, 6 ^

H

Haber, V., 62, 268, 271

Habitat, S. holbrookii, 76; B. quercicus, 112;
B. terrestris, 137, 138; A. gryllus, 161,
162, 163; P. ocularis, 211, 212; H.
andersonii, 231, 232; H. cinerea, 251, 252;
H. femoralis, 275, 276; H. gratiosa, 297;
H. squirella, 312; H. versicolor, 326; R.
aesopus, 336; R. clamitans, 355; R. grylio,
368; R. heckscheri, ^86; R. septentrionalis,
399, 400; R. sphenocephala, 422, 423; R.
virgatipes, 436-438; G. carolinensis, 456-
458

Haliburton Co., Ont., 398

Hallinan, T., 133, 318, 337, 338, 347, 348,
350, 351, 352, 384

Hallowell, E., 253, 271, 458, 475, 477

Hallowell's tree frog, 248

Hammocks, 11, 12, 13, 14, 15, 113, 122,
157, 252, 276, 280, 299, 300, 355, 426

Hammonton, N. J., 232

Hampton Co., S. C, 207, 334

Hand, H. W., 89

Hankinson, T. L., 157

Hanna, G. D., 157

Harden, Dr. J. M. B., 4, 5

Hardheads, 368

Hardwood Uplands, 162

Hargitt, C. W., 81, 106

Harlan, R., VIII, 13, 73, 74, 75, 78, 93,
105, 106, 134, 135, 138, 139, 152, 156,
161, 190, 191, 195, 200, 202, 204, 211,
276, 327, 333

Harper, F., XI, i, 3, 9, 10, 74, 89, 90, 115,
116, 117, 141, 146, 157, 170, 171, 185,
187, 188, 190, 196, 199, 200, 201, 218,
235, 237, 238, 255, 257, 277, 284, 299,
303, 315, 316, 323, 325, 328, 334, 336,
337, 338, 340, 351, 357, 371, 387, 392,
394, 402, 423, 425, 440, 441, 460, 461

Hart Lake N. Y., 397, 402, 403, 404, 405,
409, 414

Hartman, G. L., 439

Hassler, W., 3

Hastings, Fla., 4, 113, 206, 227

Hatching period, 37, S. holbrookii, 98; B.
quercicus, 128; B. terrestris, 148; A. gryl-
lus, 181; P. ocularis, 220; H. andersonii,
241; H. cinerea, 265; H. femoralis, 288;
H. gratiosa, 305; H. squirella, 320; R.
aesopus, 345; R. clamitans, 361; R. grylio,
377; R. septentrionalis, 410; R. spheno-
cephala, 430; R. virgatipes, 446; G. caro-
linensis, 467-469

Hay, O. P., 158, 165, 167, 168, 182, 183,
184, 185, 193, 195, 196, 201, 202, 203, 204

Hay, W.P., 106, 158, 165, 169, 193, 195, 253

Hebard Cypress Co., i, 9

Hebard, M., 10

Hebardsville, Ga., 316

Heckscher, A., V, i, 9

Heckscher's frog, 14, 17, 28, 31, 38, 384-397

Hellbender, 8, 9

Hellgate Ponds, N. Y., 400, 402, 405, 407

Helocaetes, 203

Helocaetes, clarkii, 198, 203, 225

486

Frogs of the Okefinokee Swamp

Helocaetes feriarum, 198, 203, 225

Helocaetes triseriatus, 198, 203, 225

Helotes Creek, Tex., 157, 205, 227

Hemidactylium scutatum, 6

Henderson, J., i, 189, 195

Hendrix, J. D., i

Henshaw, sparrow, 255

Hermit spade-foot, 73, 80, 96, 107

Hermit, spade-foot toad, 73, 87

Hermit toad, 73

Heron, 190

Helerodon platyrhimis, 190

Hickox, M., 74

Hickox, Ga., 334

High-bush blueberry, 231

Highland frogs, 138

Hill, Mr., 73

Hilliard, Florida, 13, 75, 89, 93, 94, 102,
196, 199, 201, 217, 223, 300, 309, 316,
334, 340, 343r 347, 360, 374, 375, 393

Hillsboro Co., Fla., 114

Hillsdale, Mich., 157

Hognosed snake, 152, 190

Holarctic, 66

Holbrook, J. C, VIII, IX, 4, 5, 13, 15, 63,
73, 75, 76, 77, 87, 90, 92, 95, 103, 104,
105, 106, 108, 112, 113, 114, 115, 120,
123, 125, 130, 131, 132, 133, 135, 138,
139, 140, 146, 152, 153, 154, 155, 156,
158, 168, 189, 191, 195, 196, 197, 200,
204, 205, 206, 207, 208, 209, 213, 225,
226, 227, 245, 249, 251, 255, 256, 268,
271, 272, 275, 293, 294, 312, 313, 324,
326, 327, 333, 452, 453, 454, 456, 458,

459, 475, 477

Holbrook's spade-foot, 73

Holt, E. G., 310, 324, 325, 333, 452, 477

Homestead, Fla., 74, 133, 211

Hooded warbler, 314

Hoosier frog, 397, 401

Hopkins, J. M., i, 9

Hopkins, Ga., 108, 124, 252, 280

Hop toads, 74, 133

Horned dace, 407

Horse tail, 308

Hot Springs, Ark., 206, 454

Houston, Tex., 163, 167, 248, 256, 262, 312,
314, 318, 319, 323, 326, 419, 427, 428,
453, 460, 477

Hubbard, H. G., 348

Hubbs, C. L., 162, 182, 190, 195

Hudson Bay, 43, 45, 48, 397

Hudsonian district, 155

Hunt, C. J., 439

Hurrah bush, 218, 304, 370, 392

Hurter, J., 74, 167, 195, 206, 322, 348, 435,
449, 454, 457, 474, 477

Hydrocotyle, 163, 175, 262, 386, 394

Hyla, 12, 13, 26, 29, 63, 64, 65, 69, 140,
141, 145, 168, 191, 194, 203, 225, 233,
249, 253, 272, 294, 295, 299, 301, 305,
308, 312, 322, 327, 333, 358, 460

Hyla andersonii, 15, 16, 18, 19, 21, 25, 26,
32, 33, 38, 39, 40, 53, 57, 58, 59, 60, 61,
63, 64, 65, 66, 67, 69, 158, 194, 219, 227-
248, 254, 255, 263, 271, 272, 294, 300,
310, 317, 319, 324, 435, 437, 447, 452

Hyla arborea, 65, 229, 245, 247, 452

Hyla arenicolor, 54, 65, 66, 243, 247, 294,
310, 324, 333

Hyla avivoca, 15, 65

Hyla carolinensis, 4, 62, 264

Hyla cinerea, 3, 4, 12, 13, 14, 15, 16, 18,
20, 21, 25, 26, 29, 30, 31, 32, 33, 34, 35,
37, 38, 39, 40, 54, 57, 58, 59, 60, 61, 62,

63, 64, 65, 67, 69, 163, 170, 175, 211, 229,
233, 237, 243, 246, 247, 248-271, 272,
275, 282, 286, 287, 291, 295, 299, 300,

308, 310, 316, 317, 320, 324, 358, 369,
390, 393, 466

Hyla cinerea evittata, 33, 67, 69, 229, 246,

247, 249, 255
Hyla cinerea semifasciata, 248, 253
Hyla crucifer, 34, 35, 41, 50, 51, 63, 64, 65,

66, 67, 179, 184, 194, 219, 220, 221, 225,
226, 235, 243, 309, 314, 319, 324

Hyla delitescens, 4, 5, 6

H. evittata, 249, 253, 254, 255

Hyla eximia, 65

Hyla femoralis, 3, 4, 12, 14, 15, 18, 20, 24,
25, 26, 29, 30, 31, 32, 33, 34, 37, 38, 39,
40, 55, 57, 58, 59, 60, 61, 63, 64, 65, 66,

67, 69, 92, 117, 124, 171, 175, 176, 183,
184, 194, 216, 235, 243, 244, 246, 247,
249, 261, 262, 263, 264, 265, 270, 271,
272-294, 299, 303, 305, 308, 310, 316, 318,
319, 324, 331, 333, 429, 461, 467

Hyla gratiosa, 3, 4, 12, 13, 14, 15, 17, 18,
20, 21, 22, 25, 26, 29, 31, 32, 33, 34, 35,
37, 38, 39, 40, 53, 57, 58, 59, 60, 61, 63,

64, 65, 66, 67, 69, 92, 93, 108, 113, 117,
123, 124, 129, 145, 183, 194, 215, 218,
219, 243, 246, 247, 249, 271,272,278,

282, 292, 293, 294-310, 316, 319, 320,

324,. 339

Hyla immaculata, 245

Hyla lateralis, 4, 190, 253, 313

H. miotympanuvi, 324

Hyla ocularis, 207, 225

Hyla pha,eocrypta, 65

Hyla pickeringii, 3, 7, 164, 168, 205, 225,
249

Hxjla pulchrilineata, 245, 247

Hyla regilla, 65, 66, 206, 324

Hyla semifasciata, 253

Hyla squirella, 3, 4, 5, 6, 7, 10, 12, 13, 14,
15, 18, 20, 24, 25, 26, 29, 31, 32, 34, 35,
37, 38, 39, 40, 56, 57, 58, 59, 60, 61, 63,
64, 65, 66, 67, 69, 70, 124, 183, 188, 194,
201, 219, 225, 233, 243, 244, 246, 249,
254, 255, 271, 272, 275, 277, 278, 282,

283, 286, 289, 290, 291, 293, 294, 295,

309, 310-324, 326, 331, 457, 461

Hyla versicolor, 5, 6, 13, 14, 16, 18, 20, 21,
25, 26, 29, 30, 32, 33, 34, 35, 37, 38, 39,
40, 55, 56, 57, 58, 59, 60, 61, 63, 64, 65,
66, 67, 70, 184, 235, 236, 237, 243, 247,
249, 263, 264, 271, 274, 277, 284, 285,
286, 287, 289, 290, 294, 295, 308, 310, 314,
315, 319, 324, 325-333, 360, 466

Hyla vmdis, 227, 245, 253, 271, 333

Hylidae, 28, 29, 38, 41, 50, 57, 63, 64, 65,
66, 157, 162, 177, 181, 183, 194, 229,
245, 461

Hylodes, 191, 194, 203, 225

Hylodes, crucifer, 63

Hylodes, gryllus, 63, 191, 213, 225

Index

487

Hylodes, ocularis, 204, 207, 225
Hypericum, 92, 94, 342, 443
Hypericum fasciculatum, 347, 464
Hypotheses, 65-66
Hypothetical species, 15

I
Ibises, 190, 382
Identification index, 18
Ilex cassine, 429
Ilex glabra, 116, 215, 238, 429
Ilex myrtifolia, 183, 464
Illinois, 54, I34> 156, 165, 188, 192, 195,

248, 351 » 352, 457. 477
Immaturity, 17, 18
Indian corn, 251
Indiana, 41, 56, 162, 163, 185, 188, 193,

195, 202, 204, 310, 419, 434, 452
Ingalls, O. D., 157
Inguinal amplexus, 28, 29, 30
Introduction, 1-15
Iowa, 157

Iris, 257, 262, 304, 375, 429
Island edges, 11,13
Islands, 11, 157
Ilea, 429
Ithaca, N. Y., 28, 29, 61, 153, 158, 326, 329,

330, 333, 360, 426, 433, 449

Jackson, H. H. T., 417, 419

Jackson, Ala., 272

Jackson, N, J., 245

Jackson Co., Ala., 325

Jackson Co., Ga., 140

Jacksonville, Fla., 3, 10, 113, 133, 137, 155,
156, 158, 162, 194, 196, 205, 206, 207,
262, 275, 312, 314, 316, 318, 325, 342,
365, 368, 384, 393, 419, 423, 427, 457,
459, 477

Jamaica, L. I., 95

Jamesburg, N. J., 229

Japan, 379

Jares Co., N. C, 134

Jenkins Co., Ga., 294

Jesup, Georgia, 272

Joe brown frog, 364, 371, 383

Joe pye weed, 407

Johnson Co., N. C, 454

Jones, J., 6

Jones, W. L., 5, 6

Jordan, D. S., 213

Joutel, L. H., 84

Juncus, 177

Kalm, P., 333

Kansas, 157, 163, 453, 475, 477
Keim, T. D., 450
KeUogg, R., V, 62, 68, 130
Kendall, W. C, 252, 409
Kennicott, R., 457
Kenton, Okla., 74
Kentucky, 134, 193, 195
Key West, Fla., 191
Killifish, 153, 163, 187, 212
Kinosternon subruhrum, 308
Kassimee, Fla., 365, 370
Klein, 228

(Should be Keim, T. D.)

Klots, A. B., 231, 238, 248
Klots, Mrs. E. B., 231, 238
Ivlugh, A. B., 433
Knight, H. H., 254, 255, 275

Lafayette, Miss., 453

La Grange, Ga., 453

Lake Drummond, Va., 435

Lake Ellis, N. C, 113, 132, 435, 449, 450

Lake frog, 364

Lakehurst, N. J., 84, 227, 228, 231, 236,
238, 239, 240, 243, 244, 419, 436, 437,
438, 441, 447, 448, 449, 450, 451

Lake Jesup, Fla., 454, 457

Lake Kissimee, Fla., 114, 151, 312, 323

Lakeland, Fla., 322

Lake Maxinkuckee, Ind., 188

Lake Mendota, Wise, 162

Lake Michigan, 162

Lake of Bays, Ont., 400

Lake Placid, N. Y., 397

Lake Tohopikaliga, Fla., 312

Lamar Co., Tex., 453

Land-frog, 133, 139, 152, 154

Land toad, 133

La Place, Ala., 473

Large mouth black bass, 270, 396

Larval period, 38, 39, 57, 64, S. holbrookii,
101-102; B. quercicus, 128, 129; B.
terrestris, 150; A. gryllus, 184; P. nigrita,
202; P. ocularis, 223; H. cinerea, 267;
H. femoralis, 290, 291; H. graiiosa, 307,
308; H. squirella, 321, 322; H. versicolor,
332; R. aesopus, 347; R. clamitans, 362,
363; R. grylio, 379; R. heckscheri, 394; R.
septentrionalis, 412; R. sphenocephala, 432;
R. virgatipes, 448; G. carolinensis, 472

LatreiUe, P. A., IX, 5, 14, 272, 275, 310, 352

LatreiUe's toad, 133

Laurel Hill, N. C, 133, 144

Lauren ti, J. N., 271

Lawrence, Kan., 157

Le Conte, J., IX, 4, 5, 13, 14, 78, 93, 106,
107, 108, 112, 114, 120, 131, 133, 139,
155, 156, 158, 161, 162, 164, 165, 190,
191, 193, 195, 196, 197, 199, 200, 202,
203, 204, 205, 207, 208, 213, 225, 227,
246, 251, 253, 256, 271, 272, 275, 276,
277, 293, 294, 295, 297, 304, 309, 310,
312, 313, 322, 324, 325, 327, 333, 334,
342, 344, 350, 351, 352, 453, 456, 457,

458, 460, 477
Le Cc

:)onte, J. L., 4, 6
Lee, B., i

Lee, D., i, 216, 263, 369, 379
Lee, E., i
Lee, F., i
Lee, H., i
Lee, H. H., i, 282
Lee, Jackson, i
Lee, James, 276
Lee, Joseph, i

Lee, M., i, 171, 282, 316, 328, 392, 394
Lee, Mrs., i
Lees, 113, 139, 163, 251, 270, 275, 276, 295,

359, 464, 470
Lee Co., Fla., 207, 318
Leidy, J., 228, 234, 245

488

Frogs of the Okefinokee Swamp

Leimadophis flavilatiis, io8

Leiolopisvm laterale, 252

Leister, C. W., 397, 402, 405

Lemon Citv, Fla., 314, 423, 427, 428, 458

Lenoir Co., N. C, 248

Leonard, M. D., 9, 397, 402

Leopard frog, 355

Leroy, Ala., 129, 275

Leucosomus corporalis, 407

Leucothoe, 183, 218, 304, 370, 392, 429, 443

Levy Co., Fla., 348

Liberty Co., Ga., 5, 294, 322

Liberty Co., Tex., 454

Limulus, 98

Linnaeus, 154, 271, 333

Linsdale, J. M., 163, 453, 477

Lithodytes latrans, 206

Litoria, 191

Litoria occidentalis, 205

Little Chorus frog, 13, 28, 31, 57, 206-226

Little River, Fla., 213, 314, 458

Little Sarasota Bay, Fla., 454, 474

Littleton, N. C, 460

Lizard's tail, 163, 175, 257, 282, 375, 376,
426, 428

Lloyd Co., Ga., 294

Local Okefinokee records, S. holbrookii, 74,
75; B. quercicus, 108; B. terrestris, 134;
A. gryllus, 157, 158; P. nigrita, 196; P.
ocularis, 207; H. cinerea, 248; H. fem-
oralis, 272; H. gratiosa, 294; H. squirelln,
310, 311; H. versicolor, 325; R. aesopus,
334; R. clamitans, 352; R. grylio, 365; R.
heckscheri, 385; R. sphenocephala, 419,
420; R. virgatipes. 435

Loding, H. P., 107, 113, 133, 196, 204, 297,
310, 365, 384, 419, 435, 452, 477

Loennberg, E., 109, 112, 114, 115, 133, 137,
156, 162, 165, 195, 457, 477

Logier, E. B. S., 169, 195

Long Island, 84, 85, 86, 92, 93, 95, 107,
153, 162, 194, 195, 229

Longleaf Pine Flats, 113, 294

Longleaf Pine Hills, 272, 275

Longview, Ala., 452

Lorrilliere, P., 439

Louisiana, 15, 16, 42, 44, 45, 47, 49, 50, 52,
53, 61, 65, 73, 107, 108, 113, 133, 134,
156, 174, 192, 193, 194, 195, 202, 248,
251, 254, 272, 275, 294, 312, 352, 365,
368, 369, 379, 384, 423, 450, 451, 453,

454, 457, 474
Lovendge, A., 3
Lower Austral zone, 10, 16, 74, 108, 134,

228, 228, 248, 272, 434, 453
Lower Coastal region, 10, 351
Lucania, 153
Lucania ommata, 163
Lucknow, Ont., 397, 398, 400, 414
Luxilus cornutus, 407

M
Mabee, W. B., 193, 194, 419, 434
Mabee, S. C, 158
Macclenny, Fla., 295, 334, 338
Madison Co., N. Y., 398
Madison, Wise, 162, 195
Madrid, N. Y., 397, 398
Magnolia, N. C., 107, no

Magnolia glauca, 237

Magnolia virginiana, 238, 243

Maiden-cane, 368

Maine, 56, 325, 397, 398, 402, 451

Malapoena geniculata, 339, 344

Manculus quadridigitatus, 6, 379

Manitoba, 50, 398

Mare Run, N. J., 437, 443

Marietta, Ga., 157

Martha's Vineyard, Mass., 81, 82

Maryland, 157, 163, 185, 188, 202, 245

Massachusetts, 16, 42, 73, 74, 78, 82

Mastic, L. L, 107

Mating, 26

Mating ardor, 26

Amplexation, 28; S. holbrookii, 92; B.
quercicus, 123; B. terrestris, 145; A.
gryllus, 177; P. ocularis, 218; H. ander-
sonii, 239; H. cinerea, 261, 262; H.
femoralis, 282, 284; H. gratiosa, 303; H.
squirella, 318; H. versicolor, 329; R.
aesopus, 342; R. clamitans, 358; R. grylio,
373; R. heckscheri, 389; R. septentrionalis,
406; R. sphenocephala, 426; Rana virga-
tipes, 442; G. carolinensis, 464
Coloration from Ufe 27,28;*$'. holbrookii, (^
90, 9 90; B. quercicus, cf & 9 ii9, 120;
B. terrestris, cf 143, 9 143! ^- gryllus,
cf 173, 9 173; P- nigrita, 201; P.
ocularis, 216, 217; H. andersonii, 239;
H. cinerea, 260; H. femoralis, 281; H.
gratiosa, 302; H. squirella, 317; H. versi-
color, 329; R. aesopus, 341, 342; R. clam-
itans, 357, 358; R. grylio, 372, 373; ^•
heckscheri, 388, 389; R. septentrionalis,
403, 404; R. sphenocephala, 425, 426; R.
virgatipes, 441, 442; G. carolinensis, 463
Duration, 26, 27; S. holbrookii, 91, 92; B.
quercicus, 122, 123; B. terrestris, 144; A.
gryllus, 175-177; P- ocularis, 218; H. ander-
sonii, 239; H. cinerea, 261; H. femoralis,
282, 283; H. squirella, 318; H. versicolor,
329; R. clamitans, 358; R. grylio, 373; R.
septentrionalis, 405, 406; R. sphenocephala,
426; R. virgatipes, 442; G. carolinensis, 464
Night or day, S. holbrookii, 91, 92; B.
quercicus, 122, 123; B. terrestris, 145; A.
gryllus, 175-177; P. ocularis, 218; H.
andersonii, 239; H. cinerea, 261; H.
femoralis, 282, 283; H. squirella, 318; R.
clamitans, 358; R. grylio, 373; R. septen-
trionalis, 405, 406; R. sphenocephala, 426;
R. virgatipes, 442; G. carolinensis, 464
Structural differences, 28; S. holbrookii, 90,
91; B. quercicus, 120-122; B. terrestris, 143-
144; A. gryllus, 173-175; P. nigrita, 201;
P. ocularis, 217; H. andersonii, 239; H.
cinerea, 260; H. femoralis, 281, 282; H.
gratiosa, 302, 303; H. squirella, 317-318;
H. versicolor, 329; R. aesopus, 342; R. clami-
tans, 358; R. grylio, 373; R. heckscheri, 389;
R. septentrionalis, 404, 405; R. spheno-
cephala, 425, 426; R. virgatipes, 442; G.
carolinensis, 463, 464
Mattewan, N. J., 229
Maturity, 17, 18, 63
May's Landing, N. J., 232, 234, 450
McAfee, W. L., 193, 195
Meadow frog, 27, 31, 344, 353, 359

Index

489

Meadow sweet, 407

Mearns, E. A., 114, 151, 368, 370, 398, 419

Measurements (1912-1914) 16

B. qucrcicus, iii, 112; B. terrestns, 136;
A. gryllus, 160; P. ocularis, 210; R.
grylio, 367; G. carolinensis, 455
Recent material, 16-24; S. holbrookii, 75,
76; B qucrcicus, 112; B. terrestns, 136, 137;
A. gryllus, 160, 161; P. nigrita, 198, 199;
P. ocularis, 210, 211; H. andersonii, 230,
231; H. cinerea, 250, 251; H. femoralis,
274; H. gratiosa, 296; //. squirella, 311,
312; //. versicolor, 325, 326; i?. aesopus,
335; 7?. clamitans, 354, 355; i?. (/'"^/lio, 367,
368; /?. heckscheri, 385, 386; i?. sejAen-
trionalis, 398, 399; i?. sphenoceplwla, 421,
422; /?. virgatipes, 436; G. carolineyisis,
455, 456

Medina River, Fla., 252

Meek, F. B., 474

Metiopoina fusca, 5

Aliiuulus, 407

Merrem, B., 154

Mersa Isle, Fla., 196

Mesogo7nstius, 245, 437

Metachrosis

.4. gryllus, 164, 165; P. nigrita, 199, 200;
P. ocularis, 213; i/. andersovii, 233;
i/. cinerea, 252-253; //. femoralis, 276,
277; //^. squirella, 312-314; G. caroline7isis,
458

Mexico, 46, 65, 66, 67, 324, 475

Miami, Fla., 89, 196, 211, 212. 213, 314,
323, 365

Micanopy, Fla., 309, 334, 454

Michigan, 157, 195, 352, 397

Micrnptcrus salmoides, 270

Middle Island, (L. I.), N. Y., 86

Middlesex Co., N. J., 229

Middle States, 161

Millen, Ga., 158, 248, 259, 272, 294, 301, 322

Miller, Mrs. E. P., 398, 400, 413, 414

Miller, G. S., Jr., 209, 213, 217, 224, 253,
254, 271, 398, 400, 405, 409, 413, 414, 458

Miller, W. DeW., 228, 229, 248, 435

Miller, Ind., 162

Mills Swamp, Fla., 393

Milton, Fla., 454

Mimsville, Ga., 134

Mink frog, 28, 38, 387, 419

Minnesota, 48, 56, 325, 397, 418

Mississippi, 15, 107, 133, 157, 189, 196, 199,
202, 206, 253, 254, 294, 295, 312, 364, 365

Mississippi River, 33, 41, 50, 107, 127, 153,
134, 156, 192, 193, 248, 249, 310, 434, 457,
475, 477

Missouri, 192, 195, 474, 477

Mitchell, J. D., 454

Mizzell, D., 74

Mizzell, H., I, 10, 252

Mizzell, S., I, 9

Mobile, Ala., 454

Mobile Bay, Ala., 114

Mobile Co., Ala., 113, 196, 297, 310, 365,
452, 474

Moesel, Miss J., i, 3

Moist pine barrens, 11, 13, 92, 163, 215, 224.
464

Moist woods, II, 316

Moles, 336

Monroe Co., Ind., 193, 195

Montana, 397

Montgomery, Ala., 314, 453

Montgomery Co., Ala., 310, 453

Moore, H. F., 235, 369

Moore, J. P., 228, 235, 248

Moose River, Can., 397

Morgan, T. H., 163, 195, 202, 203

Morse, M., 158, 167, 169, 195

Mt. holly, 407

Mt. maple, 407

Mt. Marcy, N. Y., 397, 409

Mt. Pleasant, S. C, 107, 134, 272, 310, 452

Mud darter, 447

Mud sunfish, 447

Myrica, 360

Munz, P. A., 62, 177, 254, 433

Musa Isle, Fla., 365, 368

Mus.Comp.vool., Harvard, 377, 244, 381, 450

Museum Essex Inst., 77

Mus. vool. Mich., 3

Myers, G. S., 33, 107, 133, 134, 156, 207,

227, 228, 248, 254, 271, 2J2, 294, 310, 324,

419, 435, 452, 460, 477
Myrica cerifera, 211
Myrica Gale, 407
Myrtle, 211

N

Narrow mouth frog, 452

Narrow mouth toad, 452, 457, 459

Narrow-mouthed frog, 452-477

Narrow-mouthed toad, 15, 28, 29, 31, 33,
38, 57, 61, 92, 359, 452-477, 476

Nash, C. W., 397

Nashville, Ga., 4, 272

Nassau Co., Fla., 10

Natrix sipedon fasciata, 382, 396, 435

Natrix sipedon sipedon, 450

Natrix taxispilata, 434

Nearctic, 66

Nebraska, 475, 477

Nebulous toad, 452

Neches River, Tex., 248

Needham, J. G., 9

Needham, J. T., 9

Nelson, G., 348

Nelson Co., Va., 189

A^eoseps, 66

Netting, M. G., 3

Neuces River, Tex., 157

Never wets, 368

New Alexandria, Va., 158, 330

New Braunfels, Tex., 157

New Brunswick, 50

New England, 16, 44, 48, 49, 73, 229, 397

New Hampshire, 397, 398

New Hanover Co., N. C., 107, 134, 207, 272

New Haven, Conn., 78, 107

New Jersey, 16, 50, 53, 74, 82, 84, 87, 106,
153, 157, 168, 186, 188, 190, 195, 227, 228,
229, 231, 232, 234, 235, 236, 238, 239, 241,
243, 244, 247, 248, 383, 419, 434, 435-452,
453

New Lisbon, N. J., 439, 443

New Madrid, Mo., 454, 457

New Madrid Co., Mo., 457

New Mexico, 157

490

Frogs of the Okefinokee Swamp

New Orleans, La., 157, 174, 192, 194, 365,

423, 474

Newport, Vt., 398, 413

New Smyrna, Fla., 348

New York, 48, 50, 73, 95, 106, 133, 155, 156,
157, 161, 195, 197, 201, 202, 203, 204, 226,
329, 333, 397, 398, 400, 402, 403, 433, 450,
451, 453, 477

Newt, 8, 215, 282, 283, 300, 415, 429, 459, 466

Nichols, Andrew, 73, 86, 89, 98, 99, 102, 103,
107

Nichols, J. T., 86, 107

Nieden, F., 134, 156, 477

Noble, G. K., 3, 33, 53, 203, 204, 206, 207,
209, 211, 225, 227, 228, 232, 233, 235, 237,
239, 240, 241, 243, 244, 245, 246, 248, 263,
435, 443, 447, 452

Noble, Mrs. R. C, 33, 53, 227, 228, 232, 233,
235, 237, 239, 240, 241, 243, 244, 245, 246,
248, 263

Norfolk, Va., 435, 460

North Carolina, 15, 42, 52, 55, 61, 93, 95,
106, 107, 108, 112, 113, 132, 133, 134, 144,
155, 156, 157, 162, 170, 177, 178, 189, 192,
193, 195, 207, 222, 226, 227, 228, 229, 237,
241, 247, 248, 254, 255, 259, 272, 294, 310,
324, 330, 419, 435, 437, 450, 451, 452, 453,
454, 460, 464, 474, 477

North Elba, N. Y., 409

Northern frog, 397

Northern leopard frog, 427

Northern meadow frog, 420, 432

Northern States, 191

Norton, A. H., 399, 402, 403

Norton, Mr., 77

Notophthalmus dor salts, 308

Notophthalmus v. viridescens, 6

Notropis chalbaeus abbottii, 437

Nuphar, 264

Nymphaea, 251, 400

Nymphoides, 262, 368

Nyssa sylvatica, 238

O
Oakdale, N. Y., 85
Oak frog, 107, 112
Oakland, Fla., 113
Oak toad, 13, 31, 38, 57, 61, 62, 107, 109, 115,

116, 117, 118, 120, 121, 122, 124, 125, 126,
127, 130, 131, 146, 212, 336, 344, 350

Ocalawacochee Slough, Fla., 318
Ocean Co., N. Y., 245
Offerman, Ga., 157
Ogcocephalus vespertilio, 340
Ohio, 195

Okefinokee locahties

Anna's pond, 12, 118, 147,150,216,300,301,

307, 309, 315, 317, 318, 319, 322
Bear House, 435, 440
Billy's Island, i, 2, 9, 10, 11, 12, 74, 108, 113,

117, 146, 150, 157, 163, 172, 184, 187, 207,
212, 213, 215, 224, 248, 251, 267, 272, 275,
276, 279, 283, 291, 295, 299, 300, 301, 309,
336, 352, 355, 358, 359, 381, 419, 424, 429,
432, 438, 458, 464

Billy Island Bay, 11, 272, 279

Billy's Lake, 11, 141, 145, 157, 163, 170, 172,
175, 187, 248, 256, 257, 258, 259, 261, 262,
264, 270, 359, 369, 371, 374, 375, 377, 470

Billy's Lake Landing, 277, 373

Black Jack Island, Ga., 108, 124, 157, 171,
190, 207, 214, 215, 216, 248, 272, 279, 424,
435, 438, 440, 448

Braganza, Ga., 13, 74, 334

Bufo pond, 141, 424

Bullfrog pond, 157

Camp CorneUa, Ga., 11, 158

Camp Island, 291

CampPinckney, Ga., 13, 14,74, 102, 108, 117,
142, 148, 150, 216, 252, 259, 272, 280, 300,
301, 312, 315, 316, 317, 318, 319, 320, 325,
326, 328, 330, 356, 359, 360, 362, 364, 387,
392, 428, 464

Cedar Hammock, 1 1

Chase Prairie, 11, 157, 162, 248

Chesser Island, i, 2, 10, 11, 12, 74, 108, 118,
126, 129, 144, 157, 158, 168, 171, 172, 173,
196, 207, 212, 215, 216, 220, 247, 248, 259,
267, 272, 280, 283, 295, 299, 300, 301, 304,
308, 309, 311, 316, 325, 329, 334, 336, 347,
352, 355, 359, 360. 380, 419, 420, 424, 427,
438, 440, 441, 443, 448, 451

Chesser School, 196, 199, 301, 307, 336, 338,

347
Chorophilus Pond, 215
Coat Bet Pond, 168, 173, 216, 279, 280, 303,

424, 425
Colerain, 216
Cornhouse Cr., Ga., 317
Crosby Pond, 116, 141, 145, 157, 212, 276
Craven's Hammock, 11, 248, 272
Cypress Pond, 157
Double-0-Bay, 11
Fargo, Ga., 108, 117, 196, 199, 200, 295, 301,

315, 384, 392
Floyd's Island, 11, 108, 157, 162, 187, 248,

272, 381
Floyd Island Bay, 1 1
Floyd's Island Prairie, 11, 157, 171, 374
Folkston, Ga., 3, 16, 74, 108, 118, 147, 158,

172, 196, 199, 207, 216, 272, 295, 300, 316,

325, 326, 329, 334, 352, 359, 364, 384, 392
GaUberry Island, 12, 157
Grand Prairie, 11, 172, 187, 248
Hickory Hammock, 11
Hog's Hole Pond, 466
Holt's Pond, 347
Honey Island, 11, 108, 113, 124, 157, 162,

207, 214, 215, 248, 272, 275, 381, 424, 458
Honey Island Prairie, 11, 113, 117, 124, 187,

212, 214, 251, 272, 420
Hurst Island, 435, 440
Indian Mound Ponds, 215, 291
Jackson Bay, 1 1

Jones Island, 12, 108, 157, 207, 215, 279
Kings Ferry, Fla., 315
Lake Sego (Seagrove), 172, 187, 259, 437
Long Pond, 141, 150, 157, 170, 171, 215, 258,

259, 261, 262, 267, 278, 282, 283, 355, 371,

373, 374, 424, 428
Middle Island, 108
Minne Island, 157

Minne Lake, 11, 163, 171, 187, 248, 267, 375
Minne Lake Run, 11, 157, 370
Mixon's Ferry, 108, 130, 157, 212

Index

491

Mixon Hammock, 11, 157, 171, 187, 219,

248, 272, 352
Moniac, Ga., 74, 108, 117, 196, 223, 295,

301, 316, 325, 326, 329, 352, 387, 393
Moonshine Pond, 141, 145, 170, 258
Murray Bay, 118, 129, 196, 223, 224
Newt Pond, 215
Nigger Pond, 347
Petty Pond, 12, 118, 172
River Narrows, 1 1
Saddlebag pond, 150, 223
Spanish Creek, Ga., 11, 142, 172, 196, 199,

316, 325, 329, 330, 355, 359
Starting Branch, 123, 142, 144, 147, 148,

158, 172, 176, 185, 187, 196, 216, 248, 252,

259, 272, 280, 295, 300, 301, 316, 317, 322,

347
Station Branch, 375
Sugar House, St. Marys, Ga., 292
Suwannee Canal, 11, 207, 212, 215, 248, 259,

264, 272, 419, 422
The Pocket, 12, 108, 117, 157, 207, 215, 248,

272, 334
Thompson Landing, 280, 316, 325, 329, 355,

359, 362, 390, 392, 393
Trader's Hill, Ga., 74, 118, 123, 129, 142,

150, 172, 207, 212, 216, 272, 2^0, 282, 299,

300, 301, 307, 309, 315, 316, 317, 321, 325,

334
Trail Ridge, 14, 334, 340, 347
Waycross, Ga., 3, 10, 93, 108, 142, 172, 248,

272, 280, 307, 316, 359, 374, 375, 440

Okefinokee region, 10

Oklahoma, 74, 107, 325. 419

Okmulgee Co., Okla., 325, 419

Old Bridge, N. J., 229

Oldfields, 1 1 , 92

Old Forge, N. Y., 397

Olney, 111., 27

Onekio, N. Y., 400, 402, 403, 405, 407, 409,

410, 414
Ontario, 169, 195, 325, 397, 398, 400, 401,

402, 405, 408, 409, 412, 414
Open lakes, 11, 12, 14, 163
Ophisaurus ventralis, 350
Opimtia, 315

Orange Co., Fla., 113, 457
Orlando, Fla., 113
Ornate chorus frog, 227
Ornate tree frog, 227
Orontium, 368, 438
Ortenburger, A. I., 74, 107, 419, 435
Osceola Co., Fla., 312
Osmunda cinnavwnea, 116
Oswego, N. Y., 397
Oswego River, N. J., 439
Otter Lake, Ont., 400, 402, 405, 409
Overton, F., 84, 85, 86, 88, 92, 93, 96, 98,

102, 107, 170, 195, 235, 333, 356, 364
Ovulation 30-32, 63, 64

Egg-laying process, S. holbrookii, 93-95;

B. quercicus, 125, 126; B. terresiris, 147;

A. gryllus, 179; P. ocularis, 219; H.

andersonii, 240; H. cinerea, 263; H.

femoralis, 285; H. gratiosa, 304; H.

squirella, 319; H. versicolor, 330; R.

aesopus, 343; R. clamitans, 360; R.

grylio, 375; R. septentrionalis, 409; R.

sphenocephala, 427; R. virgatipes, 443, 444;
Habitat, S. holbrookii, 92; B. quercicus,

123, 124; B. terresiris, 145, 146; A.
gryllus, 177; P. ocularis, 218; H. ander-
sonii, 239, 240; H. cinerea, 262; H.
femoralis, 284; H. gratiosa, 303, 304; H.
squirella, 318; H. versicolor, 330; R.
aesopus, 342; R. clamitans, 358, 359; R.
grylio, 374; R. heckscheri, 389; R.
septentrionalis, 407, 408; R. spheno-
cephala, 426, 427; R. virgatipes, 443; G.
carolinensis, 464

Humidity, 32; S. holbrookii, 93; B. querci-
cus, 125; B. terresiris, 146, 147; A. gryllus,
178, 179; P. ocularis, 219; H. cinerea, 263;
H. femoralis 285; H. gratiosa, 304; H.
squirella, 319; H . versicolor, 330; R. aeso-
pus, 343; R. clamitans, 359, 360; R. grylio,
374, 375; R. sphenocephala, 427; G. carolin-
ensis, 465
Period: S. S. holbrookii, 92-95; B. quercicus,

124, 125; B. terresiris, 146; A. gryllus,
177, 178; P. ocularis, 218, 219, H.
andersonii, 239, 240; H. cinerea, 262; H.
femoralis, 284, 285; H. gratiosa, 304; H.
squirella, 318, 319; H. versicolor, 330;
R. aesopus, 342, 343; R. clamitans, 359;
R. grylio, 374; R. heckscheri, 389, 390; R.
septentrionalis, 409; R. sphenocephala,
427; R. virgatipes, 443; G. carolinensis,
464, 465

Special: P. nigrita, 201
Temperature, 32; S. holbrookii, 93; B.
quercicus, 125; B. terresiris, 146, 147; A.
gryllus, 178, 179; P. ocularis, 219; if. cin-
erea, 263; H. femoralis, 285; H. gratiosa,
303; H. squirella, 319; H. versicolor, 330;
R. aesopus, 343; R. clamitans, 359, 360;
R. grylio, 374, 375; R. sphenocephala, 427;
G. carolinensis, 465

Ovulation without male, 30

Ozona, Fla., 108, 365

Pacific coast forms, 28, 66

Pacific coast Ranas, 28

Paine, P. R., 323

Palatka, Fla., 84

Palermo, N. J., 439

Palm Beach, Fla., 365, 368

Palmer, E. L., i, 157, 174

Palmer, Mrs. K. V. W., 157, 174, 179

Palmer, Wm., 435

Paradise Key, Fla., 322

Paris, Tex., 453, 454

Passaic River Summit, N. J., 157

Pass Christian, Miss., 248, 254, 267

Patchogue, N. Y., 84, 85, 86, 102

Paulmier, F. C, 397

Pectoral amplexus, 28, 29, 30

Pedee River, 228

Peeper, 27, 156, 168, 186, 190, 235

Pelobates cultripes, 105

Pelobates fuscus, 65, 83, 95, 452

Peltandra, 368

Pennsylvania, 82, 139, 192, 201, 203, 329,

330,' 372, 422
Pensacola, Fla., 365
Perch, 190

492

Frogs of the Okefinokee Swamp

Peromysciis gossypinus, 338, 350
Peterboro, N. Y., 398, 400, 404, 405, 409,

413, 414
Peters, J. E., 228, 234, 248
Petersburg, Va., 157, 158, 325, 453
Pickens, A. L., 134, 156, 196, 197, 204
Pickerel frog, 163, 355, 417
Pickerel weed, 257, 369, 375, 376, 387
Pickering, C, 73
Pickering's Hyla, 234
Piedmont, 228, 248, 460
Pied water snake, 434
Pierce Co., Ga., 10
Pier is phillyreifolia, 183
Pig-frog, 364, 370
Pigmy sunfish, 187, 379
Pike, N., 76, 79, 81, 87, 91, 93, 95, 100, 102,

103, 104, 107
Pilot snake, 293
Pine, G., 234, 235
Pine barrens, 11, 12, 13, 14, 15, 75, 113, 117,

163, 172, 229, 272, 297, 312, 344, 352, 443
Pinelands, S. C., 334
Pine tree frog, 272
Pine tree toad, 272, 275
Pine uplands, 162
Pine warbler, 437
Pine woods tree frog, 14, 28, 31, 92, 270,

272-294
Pine woods tree toad, 272
Pinus caribaea, 444
Pipewort, 212, 224, 438
Pirnie, M. D., 3, 10, 196, 199, 237, 292,

294. 338, 341 » 344, 351, 393, 443
Piscataquis Co., Me., 399
Pitt Co., N. C, 207
Plant habitats, 11-13
Plaquemine, La., 365
Pleasant Mills, N. J., 235
Plethodon cinereus cinereus, 6
Plethodon cinereus erythronatus, 6
Plethodon glutinosus, 4
Point Pelee, Ont., 169, 195
Poison ivy, 183, 199
Polygola cymosa, 1 83, 443
Polygonum, 163
Polygonum hydropiper, 393
Polypody, 407
Pond frog, 352
Poniederia, 145, 147, 163, 251, 257, 258, 262,

358, 373, 375, 376, 379, 387, 426
Pope, C. D., 3
Pope, E., 87
Pope, J., 476
Pope, N., 87
Pope, P. H., 162, 163, 248, 252, 256, 262,

271, 312, 314, 318, 319, 323, 324, 326, 333,

397, 399, 402, 413, 414, 419, 427, 428, 435,

453, 460, 477
Porridge Lake, Ont., 400
Portulacca oleracea, 189
Potamogeton, 401
Potamophis striatiila, 252
Potomac River, 254
Potter, D., 453, 477
Pratt, H. S., 384, 452
Prairie lake borders, 11, 12
Prairie warbler, 437

Prairies, 11, 12, 13, 14, 157, 163, 171, 172,

190, 251, 262, 270, 275, 355, 365, 375, 422
Preaxillary amplexus, 29, 30
Prickly pear, 315
Pseudacris, 7, 13, 26, 29, 63, 64, 65, 66, 68,

194, 196, 199, 201, 202, 203, 204, 205, 208.

214, 215, 218, 219, 225, 226, 294
Pseudacris clarkii, 66, 68
Pseudacris feriarum, 30, 52, 66, 68, 197, 200,

201, 203, 214, 220, 225, 226
Pseudacris nigrita, 10, 13, 15, 18, 19, 24,

25, 26, 30, 32, 33, 34, 38, 39, 40, 57, 58,

59, 60, 61, 63, 64, 66, 68, 146, 191, 192,

194, 196-204, 214, 220, 225, 235, 336, 445
Pseudacris occidentalis, 15, 69, 203, 204-206,

227
Pseudacris ocularis, 12, 13, 15, 18, 24, 25, 26,

28, 29, 31, 32, 33, 34, 37, 38, 39, 40, 52,

57, 58, 59, 60, 61, 63, 64, 65, 66, 68, 69,

161, 186, 194, 199, 203, 206-229, 278, 303,

304, 309
Pseudacris ornata, 15, 18, 19, 66, 69, 69, 203,

205, 206, 225, 226
Pseudacris septentrionalis, 68, 201, 203, 225,

226
Pseudacris triseriata 30, 33, 34, 37, 66, 68, 69,

96, 194, 197, 200, 201, 203, 209, 214, 220,

225, 226, 235
Pseudacris verrucosus, 203
Pseudobianchus striatus, 6, 10
Purslain, 189
Putnam, F. W., 95

Quarterman, H. S., i, 10

Quarterman, V. A., i, 10

Quercus catesbaei, 199

Quercus cinerea, 199

Quercus myrtifolia, 127 (should be Ilex-

myrtifolia)
Quilien, R. D., 252

R

Rabbits, 336
Rain frog, 310, 316, 452
Rainy day frog, 452, 460, 462
Raleigh, N. C, 52, 93, 106, 133, 157, 170,
178, 189, 194, 222, 272, 328, 454, 457, 464,

474, 477

Rallus virginianus, 22,^

Ramsay, E. E., 193, 195

Rana, 13, 14, 16, 2.8, 29, 63, 64, 65, 66, 85,
90, 183, 188, 191, 194, 264, 299, 315, 358,
361, 374, 387, 389, 391, 404, 409, 415, 416,
440, 457, 465, 468

Rana aesopus, 3, 4, 7, 10, 12, 14, 15, 17, 18,
20, 21, 22, 23, 25, 26, 30, 31, 32, 34, 36, 38,
39, 40, 44, 57, 58, 59, 60, 61, 63, 64, 65,
67, 70, 71, 93, 304, 308, 317, 334-352,
387, 392, 393, 418, 444, 445,

Rana agilis, 65

Rana arborea, 271, 333

Rana areolata, 7, 33, 63, 64, 65, 67, 227, 336,
342, 351, 418

Rana areolata aesopus, 350, 351

Rana areolata areolata, 4, 7, 351

Rana areolata capita, 6, 351

Rana areolata circulosa, 351

Rana cantabrigensis, 33, 417

Index

493

Rana capita, 5, 70, 334, 342, 344, 350, 351
Rana catesbeiana, 3, 4, 6, 7, 15, 16, 17, 18,

20, 21, 22, 23, 24, 25, 26, 31, 32, 33, 34, 35,
37, 38, 39, 40, 46, 47, 57, 58, 59, 60, 61, 63,
64, 66, 67, 70. 71, 237, 365, 366, 369, 370,
371, 372, 373, 376, 377, 379, 380, 381, 382,
383, 385, 387, 389, 390, 394, 395, 396, 399,
400, 412, 415, 416, 418, 441, 442, 451

Rana clairmta, 4, 84, 262

Rana clamilans, 3, 12, 13, 14, 17, 18, 20, 21,

22, 23, 25, 26, 29, 30, 31, 32, 33, 34, 35,

37, 38, 39, 40, 48, 49, 57, 58, 59, 60, 61, 63,
64, 65, 67, 70, 237, 299, 352-364, 372, 373,
376, 377, 378, 379, 380, 381, 382, 383, 385,
387, 389, 390, 393, 395, 396, 397, 399, 400,
406, 407, 411, 412, 415, 416, 417, 418, 440,
441, 442, 445, 447, 448, 449, 466

Rana cultripes Cuvier, 75, 105

Rana dorsalis, 190, 191

Rana esculenta, 65, 78, 452

Rana esculenta ridibunda, 452

Rana jontinalis, 415

Rana grylio, 3, 12, 14, 15, 17, 18, 20, 21, 22,

23, 24, 25, 26, 30, 31, 32, 33, 34, 35, 37,

38, 39, 40, 47, 48, 49, 57, 58, 59, 60, 61,

63, 64, 65, 67, 70, 71, 163, 170, 190, 257,
258, 261, 262, 267, 355, 358, 385, 387, 388,
390, 392, 396, 397, 398, 410, 411, 416, 418,
440, 441, 442, 444, 445, 447, 448, 451

Rana gryllus, 190, 202

Rana halecina, 415

Rana heckscheri, 10, 13, 14, 15, 17, 18, 20,

21, 22, 23, 25, 32, 34, 38, 39, 40, 46, 57,
58, 59, 60, 61, 63, 64, 65, 66, 67, 69, 70,
71, 266, 346, 353, 358, 383, 384-397, 448

Rana Kandiyohi, 418

Rana Le Contei, 206

Rana lentiginosa, 139, 154

Rana musica, 154

Rana nigrita, 196, 202

Rana onca, 47, 442, 450

Rana palustris, 34, 36, 44, 45, 63, 64, 334,

350, 351, 401, 403, 407, 409, 417, 418, 445
Rana pipiens, 3, 5, 6, 7, 15, 34, 36, 45, 46, 63,

64, 65, 67, 71, 334, 340, 344, 347, 348, 350,

351, 402, 418, 419, 420, 421, 422, 423, 426,
427, 428, 430, 431, 433, 434, 443, 444, 445,
450

Rana septentrionalis, 15, 17, 18, 20, 21, 22,
23, 25, 26, 29, 32, 36, 37, 38, 39, 40, 48,
57, 58, 59, 60, 61, 63, 64, 65, 67, 70, 71,
350, 379, 383, 394, 397-419, 438, 444,
448, 451

Rana sphenocephala, 3, 7, 12, 13, 14, 17, 18,
20, 21, 22, 23, 25, 26, 27, 29, 30, 31, 32, 33,
34, 36, 37, 38, 39, 40, 45, 57, 58, 59, 60,
61, 63, 64, 65, 67, 71, 146, 175, 229, 251,
290, 340, 341, 343, 344, 347, 348, 350, 351,'
373, 379, 418, 419-435, 441, 444, 445, 470

Rana ST/Ivatica, 33, 36, 65, 66, 344, 417, 444,
445

Rana temporaria, 451

Rana temporaria pretiosa, 416

Rarm terresiris, 154

Rana virescens sphenocephala, 4, 434

Rana virescens virescens, 4, 6

Rana virgatipes, 10, 12, 14, 16, 17, 18, 20, 21,
22, 23, 25, 26, 30, 31, 32, 33, 34, 36, 37,
38, 39, 40, 48, 49, 50, 57, 58, 59, 60, 61,
63, 64, 65, 66, 67, 69, 71, 158, 183, 194,
229, 237, 238, 240, 272, 299, 346, 350,
373, 381, 383, 392, 398, 401, 407, 409, 415,
418, 435-452

Rancocas Cr., N. J., 439, 443

Range 15-16, S. holbrookii, 73-75; B. quercicus
107, 108; B. terresiris, 133, 134; A.
gryllus, 156-158; P. nigrita, 196; P.
ocularis, 206, 207; H. andersonii, 228, 229;
H. cinerea, 248; H. femoralis, 272; H.
gratiosa, 294; H. squirella, 310, 311; H.
versicolor, 325; R. aesopus, 334; R.
clamitans, 352; R. grylio, 364, 365; R.
heckscheri, 384, 385; R. septentrionalis,
397, 398; R- sphenocephala, 419, 420;
R. virgatipes, 435; G. carolinensis, 452-454

Ranidae, 28, 33, 41, 44, 58, 63, 66, 158, 183.

324, 477
Ranunculus, 401, 406
Raritan river, N. J., 229
Rattler, 156, 166
Red bamboo, 163
Red-bellied dace, 407
Red-bellied horn snake, 457
Red-headed scorpion, 300
Red maple, 407
Red toads, 138, 139
Reed, H. D., 228
Reese, A. M., 323
Refugio Co., Tex., 476
Regal's fern, 407

Rehn, J. A. G., 10, 84, 228, 245, 439
Reynolds, A. G., 348
Rhexia, 92, 94
Rhexia mariana, 443, 444
Rhineura, 66

Rhinichthys atronasus, 407
Rhoads, S. N., 155, 156, 193, 195, 457, 477
Rhus, 199
Rhus Vernix, 238
Ribbon snake, 224, 270, 309, 434
Rice, F. L., 351
Riceboro, Ga., 4, 5, 6, 134, 204, 205, 272,

334, 350, 351, 454
Richardson, J., 197
Richland Co., 111., 162, 195
Richmond, Va., 453
Richmond Co., N. C, 248
Rider, W. R., 314, 315
Ridgway, Robert, 368, 370
Ridgway Bog, Wise, 417
Ringneck snake, 379
River swamp frog, 384
River swamps, 11, 13, 14, 355, 422, 426
Roadsides, railroads, around buildings, 1 1 ,

12, 13, 14, 422
Roan Mt., Tenn., 155
Roanoke River, N. C, 330
Rockies, 47

Rockingham, N. C, 228, 248
Rocky Lake, Fla., 21 r
Rocky Mountain frog, 397, 401
Rogers, J. T., 323, 324
Roswell, N. M., 157
Rough chorus frog, 196

494

Frogs of the Okefinokee Swamp

Royal Palm Hammock, Fla., 133, 211, 310,

365, 370, 419, 452
Runyon Pond, N. J., 229
Ruthven, A. G., 3, 169, 178, 189, 195
Ryder, J. A., 460, 464, 465, 467

S
Sahal serenoa, 314
Sabalian zone, 10, 15, 16, 65, 66, 69, 108,

207, 226, 272, 309, 365
Sabbatia decandra, 183
Sackett, R., 228

Sackett's Harbor, N. Y., 413, 414
Sagiitaria, 147, 386, 394
St. Clair Co., Ala., 452
St. George, Ga., 280, 301, 307, 317, 359, 360,

374, 375

St. John's wort, 342

St. Louis, Mo., 195, 457, 477

St. Louis Co., Mo., 457

St. Mary's, Ga., 108, 118, 142, 172, 216, 259,
272, 292, 316

St. Mary's River, 10, 11, 12, 14, 16, 108,
117, 158, 171, 216, 248, 252, 272, 279, 280,
295, 311, 315, 316, 325, 326, 334. 338, 347,
352, 355, 356, 359, 362, 364, 384, 387, 389,
390, 392, 393, 420, 464

St. Petersburg, Fla., no, 113, 125

St. Simon's Island, Ga., 4, 5

St. Tammany Parish, La., 108, 454

Salamandra auriculata 5

Salamandra hilineata, 5

Salamandra erythronota, 5

Salamandra fasdata, 5

Salamandra guttolineata, 5

Salamandra quadridigitata, 5

Salamandra quadrimaculata, 5

Salamandra sabnonea, 5

Salamandra talpoidea, 5

Salamandra venenosa, 5

Salem, Mass., 73

San Antonio, Tex., 61, 157, 252, 418

San Francisco, Cal., 205

Sand bluffs, 1 1

Sand Mt., Ala., 325

Sand ridges (hills), 11, 13, 336, 352

Sand scrub, 1 1

Santo Domingo, 245

Sandwich Islands, 131

Saranac Lake, N. Y., 402

Sarracenia minor, 212

Satilla River,. Ga., 157

Saururus cernuus, 175, 257, 258, 262, 282,

375, 376, 426, 428
Savanna cricket, 156, 158
Savannah, Ga., 384, 392, 394
Savannah cricket, 156, 161, 202
Savannah cricket frog, 156, 168, 191, 194
Savannah River, 228, 384

Saw palmetto, 212

Say, T., 155

Sayreville, N. J., 229

Scaphiopodidae, 41, 57

Scaphiopus, 13, 26, 30, 65, 85, 87, 93, 95,

97, 100, 240, 465, 467
Scaphiopus couchi, 66, 68, 85, 91, 103, 106
Scaphiopus hammondi, 66, 68, 85, 91, 97,

106

Scaphiopus holbrookii, 6, 10, 13, 16, 17, 18,
20, 22, 25, 26, 29. 32, 33, 34, 36, 37, 38, 39,
40, 42, 57, 58, 59, 60, 61, 65, 66, 68, 73-
107, 109, 146, 229, 240, 304, 336, 445, 452,
464, 472

Scaphiopus holbrookii albus, 15

Scaphiopus hurteri, 66

Scaphiopus rectifrenis, 106

Scaphiopus solitarius, 5, 78, 80, 83, 95, 105,
106, 107

Sceloporus spinosus, 252

Sceloporus variabilis, 252

Schmidt, K. P., 107, no, 133, 134, 156, 228,
248, 271, 272, 294, 310, 324, 452, 477

Schneider, J. G. IX, 14, 268

Schultz, Miss E. D., 252

Schwartz, E. A., 322

Scotland Co., N. C, 133, 134, 144

Scraper Frog, 272, 310, 315, 316

Screaming Frog, 352

Screven, Ga., 248, 272, 295, 301, 453

Sebastian, Fla., 348

Semoiilus atromaculatus, 407

Seventh Lake, N. Y., 400

Shackleford's Banks, N. C, 113

Shad frog, 419, 422,

Shaw, George, 134, 137, 138, 139, 152, 155,
156

Shelby Co., Ala., 453

Shelf ord, V. E., 417

Sherman, F. Jr., 113. 228

Sherwood, W. L., 95, 98, 102, 107, 167, 177,

195
Shinn, E., 448
Shinn, W., 448
Shortleaf Pine Hills, 162
Shortlined skink, 252
Shrews, 336

Shufeldt, R. W., 174, 338, 340, 349, 352
Shute, J. G., 450

Sibley, C. K., 397, 402, 403, 405, 414
Sierra Nevada, 46
Sigmodon, 338
Simpson, S. E. R., 401, 405, 407, 408, 409,

413
Single eggs, 31, 35, 37, 64
Siren intermedia, 5
Siren lacertina, 4, 10, 429
Siren Run, 212, 429, 430
Siren striata, 5
Six-hned hzard, 350
Skink, 379
Skinner, H., 229
Small loose egg mass, 30, 37
Smilax, 116, 212, 231, 237, 371, 429, 437
Smilax glauca, 231
Smilax walteri, 443
Smith, F. S., 78, 107
Smith, H. M., 213
Smith ville, N. C, 107
Smooth chorus frog, 204, 206
Snyder, C. E., 334
SoUtary spade-foot, 28, 31, 38, 39, 57, 61,

73, 75
Sohtary toad, 73
Soudan, Va., 157
South America, 105
South Atlantic States, 193

Index

495

South Carolina, 15, 44, 46, 50, 52, 53, 65,
73, 78, 86, 92,93, 107, 112, 133, 134, 138,
139, 156, 157, 158, 196, 197, 202, 204, 205,
206, 207, 208, 227, 228, 237, 241, 248, 259,
272, 294, 310, 323, 324, 334, 351, 365, 435,
437, 452, 453, 456, 475, 477

South River, N. J., 229

Southern bullfrog, 14, 17, 28, 31, 38, 176,

258. 364, 387
Southern gopher frog, 334
Southern leopard frog, 419-435
Southern meadow frog, 14, 28, 30, 39, 58,

141, 419-435
Southern parula warbler, 314
Southern Pines, N. C, 228, 240, 435
Southern riband snake, 190, 292, 323
Southern States, 161, 245, 419
Southern tickseed, 163
Southern toad, 13, 30, 38, 57, 61, 123, 126,

131, 133-156, 344
Southern tree frog, 14, 28, 31, 124, 310-324.
Southern tree toad, 310
Southern water snake, 153, 382, 396, 435
Southport, N. C, 107
Spade-foot, 13, 28, 29, 30, 33, 73, 74, 77,

78, 79, 80, 82, 88, 91, 92, 93, 94, 95, 97,

98, 100, loi, 102, 104, 105, 106, 280, 303,

317, 336, 457
Spade-foot toad, 73, 83, 84, 85, 86, 89, 93,

106, 107
Spanish curlews, 300
Spatterdocks, 400
Spawning order, 57
Spawning period, 32, 33, 35-37, 38, 63
Spea, 106

Spea hammondi, 106
Spea multiplicata, 106
Species accounts, 73-477
Speedwell, N. J., 439
Spelerpes bilineatus 6
Spelerpes bislineatus, 6
Spelerpes guttolineatus, 6
Spelerpes ruber stictoceps, 6
Speonk, N. Y., 85
Sphagnum bogs, 11, 426
Sphagnum frog, 240, 383, 392, 435-45-?
Spice bush, 339
Spiny swift, 252
Spooner, C. S., 9
Spotted frog, 419, 420
Spreading adder, 131
Spring Cr., Ga., 157, 262
Springfield, Mass., 76, 77, 106
Springfield Nat. Hist. Museum, 77
Spring frog, 187, 352, 419, 423, 427, 432
Squirrel Hyla, 310
Squirrel tree frog, 310, 314
Squirrel tree-toad, 310
Stafford's Forge, N. J., 245
Staked Plains, Tex., 156
Staten Island, 84
Steiner, G., 270
Stejneger, Leonhard, III, V, IX, 14, 73, 107,

133, 196, 203, 227, 248, 255, 271, 272, 294,

310, 325, 334, 352, 364, 365, 366, 368,
^ 370, 381, 382, 384, 397. 419, 435, 452, 476
Stereochilus marginatus, 6
Sternotherus minor, 396
Stone, W., 156, 195, 228, 232, 248, 438, 452

Storer, D. H., 5

Storer, T. I., V

Storeria dekayi, 252

Storm frogs, 73, 74, 75

Storm toads, 73 75

Strands 11, 12, 157, 163, 215

Strecker, J. K. Jr., V, 74, 156, 162, 177, 178,

195, 206, 453, 454, 459, 464, 476, 477
Striped tree-frog, 196
Structural characters (1912), B. quercicus,

no, in; B. terrestris, 135; A. gryllus,

159; P. nigrita, 198; P. ocularis, 210; H

andersonii, 230; H. cinerea, 250; H. femor-

alis, 274; H. gratiosa, 295, 296; R. clami-

tans, 353, 354; R. grijlio, 366, 367; R.

sphenocephala, 421; G. carolinensis, 455
Sudden appearance and disappearance of

spadefoots, 73
Suggestions, 67-71
Supplemental records, S. holbrookii, 73, 74;

B. quercicus, 107, 108; B. terrestris, 133,

134; A. gryllus, 156, 157; P. nigrita, 196;

P. ocularis, 206, 207; H. andersonii, 228,

229; H. cinerea, 248; H. femoralis, 272;

H. gratiosa, 294, 295; H. squirella, 310;

H. versicolor, 325; R. aesopus, 334; R.

clamitans, 352; R. grylio, 365; R. heck-

scheri, 384; R. septentrionalis, 397, 398;

R. sphenocephala, 419; R. virgatipes, 435;

G. carolinensis, 452-454
Supraxillary amplexus, 28, 29, 30
Sm-face, H. A., 433
Surface egg films, 31, 33- 35, 37, 64
Surinam, 154
Suwannee River, 11, 108, 130, 171, 204, 248,

272,311,315,370
Swamp bullfrog, 369, 370
Swamp chorus frog, 196
Swamp cricket frog, 57, 196, 202, 206, 336
Swamp tree frog, 169, 196, 206
Swamp tree-toad, 196
Swinnerton, A., 87
Swinnerton, J., 87

T
Tadpoles, 39-56, 64, Color description from

life, S. holbrookii, 98; B. quercicus, 128;

B. terrestris, 148; A. gryllus, 181; P.

ocularis, 220, 221; H. andersonii, 241; //.

cinerea, 265; H. femoralis, 288; H.

gratiosa, 306; H. squirella, 320; //.

versicolor, 330, 331; R. aesopus, 345; R.

clamitans, 361; R. grylio, 377; R. heck-

scheri, 390, 391; R. septentrionalis, 410; R.

sphenocephala, 430; R. virgatipes, 446;

G. carolinensis, 469
General appearance, 98-99, S. holbrookii,

98-99; B. terrestris, 148, 149; A. gryllus,

181; P. ocularis, 221; H. andersonii, 241,

242; H. cinerea, 265; H. femoralis, 288;

H. gratiosa, 306; H. squirella, 320; H.

versicolor, 331; R. aesopus, 345, 346; R.

clamitans, 361; R. grylio, 377, 378; R.

heckscheri, 391; R. septentrionalis, 411; R.

sphenocephala, 430, 431 ; Gr. carolinensis, 469
General remarks, S. holbrookii, 99-101; B.

quercicus, 128; B. terrestris, 150; A.

gryllus, 182-184; P. nigrita, 202; P.

ocularis, 222, 223; H. andersonii, 243; H.

cinerea, 266, 267; H. femoralis, 289, 290;

496

Frogs of the Okefinokee Swamp

H. gratiosa, 307; H. sqidrella, 321; H.

versicolor, 332; R. aesopus, 346, 347; R.

damiians, 362; R. grylio, 378, 379; R.

heckscheri, 392-394; R. sphenocephala, 432;

R. virgatipes, 447; G. carolinensis, 470, 471
Measurements, S. holbrookii, 99; B.

terrestris, 149, 150; A. gryllus, 182; P.

ocularis, 222; H. andersonii, 242; H.

cinerea, 266; i7. femoralis, 289; //.

gratiosa, 306, 307; ^. squirella, 321; //.

versicolor, 331, 332; i?. aesopus, 346; i2.

clamitans, 362; ^. grylio, 378; E. /iecA>

scheri, 391, 392; K. septentrionalis, 411,

412; ij. sphenocephala, 431; i?. virgatipes,

447; G. carolinensis, 470
Mouth parts, 40, 41-56, S. holbrookii, 99;

B. terrestris, 149; A. grijllus, 181, 182; P.

ocularis, 221, 222, i/. andersonii, 242; i/.

cinerea, 266; i/. femoralis, 288, 289; //.

gratiosa, 306; /f. squirella, 320, 321; i/.

versicolor, 331; -R. aesopus, 346; /?.

clamitans, 361, 362; i2. grylio, 378; /?.

heckscheri, 391; R. septentrionalis, 411; 72.

sphenocephala, 431; jR. virgatipes, 446;

(j. carolinensis, 470
Tall meadow rue, 407
Tallulah Falls, Ga., 329
Tantilla coronaia, 108
Tantilla nigriceps, 252
Tappahannuk, Va., 89
Taylor, W. J., 4, 204
Teimatobius, 139

Temiessee, 73, i55, 156, I93, I95, 457, 477
Texas, 15, 16, 33, 41, 42, 44, 50, 54, 55, 56,

61, 134, 154, 156, 157, 162, 163, 178, 193,

194, 195, 201, 202, 203, 204, 205, 206,

226, 227, 229, 248, 252, 255, 256, 263,

272, 310, 312, 319, 323, 324, 325, 326,

365, 418, 419, 427, 434, 452, 453, 454,

458, 476, 477
Thaninophis sackern, 190, 224, 292, 434
Thamnophis sirtalis, 292, 475
The barker, 294, 300, 301
The cricket Hylodes, 156
The northern tree-toad, 325
Theodore. Ala., 157, 365, 371, 375
The scraper, 312, 315, 316
Thompson, Miss C, 169, 178, 189
Thompson, Mi.'is H., 169, 178, 189
Thompson, R. J., 114
Throats in males, 28
Thumb excrescences, 28
Tifton, Ga., 196
Toads, 30, 33, 57, 100, 336, 359
TomlMgbee River, 272
Toothed haw, 407
Toothless frog, 452, 454
Topsfield, Mass., 73
Towhee, 437
Townsend, J. H., 131
Transformation 17, 18, 56-59, 65,

General remarks, A. gryllus, 186-188; P.

nigrita, 202; H. femoralis, 291, 292; H.

gratiosa, 308, 309; H. squirella, 322; R.

clamitans, 363; R. grylio, 380, 381; R.

sphenocephala, 433; R. virgatipes, 449; G.

carolinensis, 473
Period, 56, 57, 64; S. holbrookii, 102; B.

quercicus, 129; B. terrestris, 150; A.

gryllus, 185; P. ocularis, 223; H. ander-
sonii, 243, 244; H. cinerea, 267; H.
femoralis, 291; H. gratiosa, 308; H.
squirella, 322; H. versicolor, 332; R.
aesopus, 347; R. clamitans, 363; R. grylio,
379. 380; R. heckscheri, 394, 395; R. sep-
tentrionalis, 412-414; R. sphenocephala,
432; R. virgatipes, 448; G. carolinensis, 472
Size, 38, 39, 40, 59, 64; S. holbrookii, 102;
B. quercicus, 129; B. terrestris, 150, 151;
A. gryllus, 185, 186; P. ocularis, 223, 224;
H. andersonii, 244; H. cinerea, 267; H.
femoralis, 291; H. gratiosa, 308; H.
squirella, 322; H. versicolor, 332; R.
aesopus, 347, 348; R. clamitans, 363; R.
grylio, 380; R. heckscheri, 395; R. septen-
trionalis, 412-414; R. sphenocephala, 432;
R. virgatipes, 448, 449; G. carolinensis,
472, 473

Transition zone, 66

Tree frogs, 33, 38, 57, 61, 65, 248

Tree toad, 14, 176

Trenton, N. J., 87, 88, 234

Triton niger, 5

Triturus viridescens dorsalis, 10

Troost, G., 73

Troy, J., XI

Tufted titmouse, 314

Turkey buzzard, 437

Turkey Lake, Ind., 162, 194

Turtles, 153, 190, 293, 396

Tuscaloosa Co., Ala., 452

Tuskegee, Ala., 314, 453

Tympana in males, 28

Type locality, S. holbrookii, 73; B. quercicus,
107; B. terrestris, 133; P. nigrita, 196; P.
ocularis, 206; H. andersonii, 228; H.
cinerea, 248; H. femoralis, 2^2; H. grati-
osa, 294; H. squirella, 310; R. aesopus,
334; R. grylio, 364; R. heckscheri, 384;
R. septentrionalis, 397; R. virgatipes, 435;
G. carolinensis, 452

U

Uhler, Dr., 245

Umbra, 245, 447

United States, 73, 107, 132, 133, 134, 166,
181, 192, 193, 205, 209, 240, 245, 305,
333, 382, 386, 391, 475, 476, 477

U. S. Bureau of Fisheries, 369, 392, 394,
398, 407

U. S. Nat. Museum, 103, 113, 114, 120,
125, 138, 151, 153, 174, 188, 203, 206,
209, 213, 217, 224, 253, 309, 312, 318,
322, 323, 324, 348, 349, 350, 381, 398,
400, 405, 409, 413, 414, 415, 449, 457,
458, 474, 475, 476

Upper Austral zone, 66, 157, 226, 228, 248

Upper Coastal region, 10, 228

Upton, G. B., 254, 397, 402, 405

Utricularia, 264, 368, 444

V

Vaccinium atrococcum, 238, 243
Vaccinium corymbosum, 238, 243
Vaccinium myrsinites, 277
Van Buren, Me., 399

Van Hyning, O. C, 196, 199, 200, 213, 227,
384, 390

Index

497

Van Hyning, T., 74, 93, 107, 297, 304, 310,
337- 343, 352

Variable swift, 252

Variation in color, B. quercicus, 114; B.
terrestris, 138, 139; A. gryllus, 165, 166;
P. nigrita, 200; P. ocularis, 213; H.
andersonii, 233; H. cinerea, 253-255; G.
carolinensis, 458, 459

Vermont, 397, 398

Viburnum nudum, 238

Victoria, Tex., 454, 476

Viereck, H. L., 232

Vigo Co., Ind., 163, 194

Village streets, 1 1

Viosca, P., Jr., 44, 52, 53, 61, 107, 108,
113, I33> 134, 156, 157, 162, 193- 194,
195, 251, 271, 272, 275, 294, 310, 312,
324, 365, 368, 369, 370, 379, 383, 384,
419, 423, 435, 450, 457

Virginia, 15, 16, 41, 54, 56, 89, 106, 134,
152, 157, 158, 176, 177, 189, 229, 248,
255, 310, 325, 330, 372, 435, 452, 453,
460, 477

Virginia rail, 234

Virginia willow, 163

Vitelli, 25, 34-37, 38, 39

Vitelline membranes, 34, 35

Vocal sacs, 26, 63

Voice, 26, 63; S. holbrookii, 74, 86-90; B.
quercicus, 11 5-1 19; B. terrestris, 140-143;
A. gryllus, 168-173; P- nigrita, 200, 201;
P. ocularis, 214, 216; H. andersonii, 234-
238; H. cinerea, 256-259; H. femoralis,
277-281; H. gratiosa, 298-301; H. squir-
ella, 314-317; H. versicolor, 327-329;
R. aesopus, 339-341; R- clamitans, 355-
357; R. grylio, 370-372; R. heckscheri,
387, 388; R. septentrionalis, 402, 403;
R. sphenocephala, 423-425; R. virgatipes,
438-441; G. carolinensis, 459-463

1921 record; B. quercicus, 117; B. terres-
tris, 141, 142; A. gryllus, 170, 171; P.
ocularis, 215, 216; H. cinerea, 258, 259;
H. femoralis, 278, 279; H. gratiosa, 300,
301; H. squirella, 314, 315; H. versicolor,
329; R. clamitans, 356; R. grylio, 371;
R. sphenocephala, 424; R. virgatipes, 440,
441; G. carolinensis, 461, 462

1922 record; B. quercicus, 118; B. terres-
tris, 142; A. gryllus, 172; P. ocularis, 216;
H. cinerea, 259; H. femoralis, 280, 281;
H. gratiosa, 301; H. squirella, 315, 316;
H. versicolor, 329; R. clamitans, 356; R.
sphenocephaly, 424, 425; R. virgatipes, 441 ;
G. carolinensis, 462

W

Wabash Valley, 27

Waco, Texas, 178, 206

Wading River, N. J., 439

Wake Co., N. C, 454

Waldo, Fla., 384

Walker, Mrs. J. L., 2

Wampee, 368

Ware Co., Ga., 10

Washington, D. C, 248, 253, 272

Washington Co., Ala., 272

Washington Parish, La., 108

Water courses, 11, 14, 365, 375

Water frog, 419

Water hyacinth, 386, 394

Water lily, 175, 251, 264, 368, 386, 394,

400, 407, 438
Water-lizards, 8
Water moccasin, 434, 457
Water pennywort, 163, 175, 386
Water-shields, 368

Water snake, 270, 309, 361, 415, 450
Wayne Co., Ga., 295
Wayne Co., N. C, 454
Waynesville, Ga., 272, 276, 292
Weber, J. A., 203, 206, 209, 211, 213, 318
Weed, A. C, 418, 419
West, L., 397, 402
West Indies, 15
West Palm Beach, Fla., 348
West Point, Ala., 453
Western cricket, 156
Western cricket frog, 156
Weymouth, N. J., 238
Wheeler, W. M., 84
White, G., 5

White eyed vireo, 314, 437
White Lake, N. Y., 397, 400
Whiting, N. J., 439
Wilce Co., Ala., 248
Wilder, B. G., 228, 244
Williams, J. L., 138, 156
Wilmington, N. C, 107, 133, 134, 156, 207,

229, 248, 254, 272, 294, 365, 384, 452,

460, 474, 477
Wilson, W. J., Mr. and Mrs., i
Winona Lake, Ind., 195
Wintering tadpoles, 38, 39, 46-50, 57-58, 64
Wisconsin, 162, 195
Wittfield, W., 323
Wood, N. R., 114, 337, 348
Wooded watercourses, 11, 270
Wooded lake edges, 1 1 , 365
Wood frog, 205
Woodpecker, 355
Woods Hole, Mass., 153
Woodicardia 124, 218, 2:
„443

Worsham, E. L., 9
Wray, Colo., 157
Wrens, Ga., 157
Wright. Mrs. A. A., V, XI,

96, 107, 126, 127, 133,

180, 196, 220, 227 238,

264, 271, 284, 310, 324,

351, 374, 375, 384, 387,

428, 435, 441, 445, 447,
„ 471, 477
Wright, H. P., 33
Wurdemann, G., 474

X

Xyris, 94, 368, 443, 444

Y

Yaphank, N. Y., 86
Yar ow, H. C, 155. 156
Yellow-throated Green Frog, 352
Yellow water lily, 264

Z

Zea mais, 251

3, 304, 368, 440,

3, 9,
146,

243,
330,
397,
448,

o, 74, 94,
148, 156,
252, 262,

337, 341,
407, 409,
452, 467,