• TABLE OF CONTENTS
HIDE
 Front Cover
 Front Matter
 Title Page
 Table of Contents
 List of Illustrations
 Amphibians and land reptiles of...
 The fishes of Porto Rico and the...
 The fishes of Porto Rico and the...
 The Ascidians of Porto Rico and...
 Index for volume X: Amphibians...
 Plates
 Back Matter
 Back Cover














Title: Scientific survey of Porto Rico and the Virgin Islands
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Permanent Link: http://ufdc.ufl.edu/UF00091487/00012
 Material Information
Title: Scientific survey of Porto Rico and the Virgin Islands
Alternate Title: Scientific survey of Puerto Rico and the Virgin Islands
Physical Description: 19 v. : ill. ; 24 cm.
Language: English
Creator: New York Academy of Sciences
Jay I. Kislak Reference Collection (Library of Congress)
Publisher: The Academy,
The Academy
Place of Publication: New York N.Y
Publication Date: 1928-1930
Frequency: completely irregular
 Subjects
Subject: Scientific expeditions -- Periodicals   ( lcsh )
Natural history -- Periodicals -- Puerto Rico   ( lcsh )
Natural history -- Periodicals -- Virgin Islands of the United States   ( lcsh )
Natuurlijke historie   ( gtt )
Geologie   ( gtt )
Expedities   ( gtt )
Genre: bibliography   ( marcgt )
Spatial Coverage: Puerto Rico
United States Virgin Islands
 Notes
Summary: Includes bibliographies.
Ownership: Provenance: Gift of Jay I. Kislak Foundation.
Statement of Responsibility: New York Academy of Sciences.
Dates or Sequential Designation: Vol. 1, pt. 1-
Dates or Sequential Designation: Ceased with vol. XIX, pt. 1.
General Note: Latest issue consulted: Vol. 18, pt. 4 (1952).
General Note: Kislak Ref. Collection: Vol. 18, pt. 2 (1941)-pt. 4 (1952).
 Record Information
Bibliographic ID: UF00091487
Volume ID: VID00012
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 01760019
lccn - 2002209050

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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
    Front Matter
        Front Matter 1
        Front Matter 2
    Title Page
        Page i
        Page ii
    Table of Contents
        Page iii
    List of Illustrations
        Page iv
        Page v
        Page vi
        Page vii
        Page viii
        Page ix
        Page x
        Page xi
        Page xii
        Page xiii
        Page xiv
        Page xv
        Page xvi
    Amphibians and land reptiles of Port Rico with a list of those reported from the Virgin Islands
        Page 1
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    The fishes of Porto Rico and the Virgin Islands: Branchiostomidae to Sciaenidae
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    The fishes of Porto Rico and the Virgin Islands: Pomacentridae to Ogcocephalidae
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    The Ascidians of Porto Rico and the Virgin islands
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    Index for volume X: Amphibians and land reptiles
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    Plates
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    Back Matter
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    Back Cover
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Full Text




















_-- ru





,0___
l^ -







NEW YORK ACADEMY OF SCIENCES


SCIENTIFIC SURVEY
OF

Porto Rico and the Virgin Islands


VOLUME X


NEW YORK:
PUBLISHED BY THE ACADEMY
1930






















CONTENTS OF VOLUME X
Page
Title-page........................................................ i
C ontents ........................ ............................. ........ iii
D ates of Publication of Parts............................................ iii
List of Illustrations ...................... .... ....... .................. iv
Amphibians and Land Reptiles of Porto Rico, with a List of Those Reported
from the Virgin Islands. By KARL PATTERSON SCHMIDT. ............... 1
The Fishes of Porto Rico and the Virgin Islands-Branchiostomidae to Sciae-
nidae. By J. T. NICHOLS.......................... ................... 161
The Fishes of Porto Rico and the Virgin Islands-Pomacentridae to Ogcoce-
phalidae. By. J. T. NICHOLS ....................................... 297
The Ascidians of Porto Rico and the Virgin Islands. By WILLARD G. VAN
NAME ............................................................... 401
Index ...................... ........................ ............... 513


DATES OF PUBLICATION OF PARTS

Part 1, November 22, 1928.
Part 2, September 10, 1929. 7 7 2.
Part 3, March 15, 1930
Part 4, August 1, 1930












(iii)







N 'IENTIFIC ,P' 'EY OF PORTO RICO


LIST OF ILLUSTRATIONS

FIGURES
Page
Head of toad (left) contrasted with Leptodactylus (right).................... 30
Foot of Leptodactylus (left) contrasted with foot of Eleutherodactylus (right).
Compare slender and expanded tips of digits .......................... 30
Head of Bufo lemur................................................. 32
Habitus of juvenile Bufo lemur (A), with side view of head (B). A. M. N. H.
No. 10151. Natural size ........................................... 33
Bufo m arinus. M ayagiiez .............................................. 35
Leptodactylus albilabris, to show variation in color pattern and in form of snout.
A. M. N. H. No. 10125 (A and B); A. M. N. H. No. 10143 (C and D).
Natural size ......................... ......................... 39
Lep'o lactylus dominicensis. A. M. N. H. No. 20952 for comparison with
L. albilabris. Three-fourths natural size.... ........ ...... ... 39
Lateral view of tadpole of Leptodactylus albilabris .......................... 42
Mouth parts of tadpole of Leptodactylus albilabris .......................... 43
Inside of mouth of Eleutherodactylus portoricensis (left) and of E. richmondi
(right), showing different arrangement of the vomerine teeth. ........... 45
Three common color variants of Eleutherodactylus portoricensis. A, A. M. N. H.
No. 10139; B, No. 10243; and C, No. 10249. Natural size.............. 46
Embryo of Eleutherodactylus portoricensis. A. M. N. H. No. 10302. Six
tim es natural size.................................... ........... 48
Peter's figures of the embryo of Eleutherodactylus cortoricensis............... 49
Elcutherodactylus gryllus. A. M. N. H. No. 10226. Twice natural size. ..... 51
Eleutherodactylus locustus. A. M. N. H. No. 10240. Twice natural size...... 53
Eleutherodactylus cramploni. A. M. N. H. No. 10305. Twice natural size. ... 55
Eleutherodactylus antillensis. A. M. N. H. No. 10019. Twice natural size.... 57
Eleutherodactylus brittoni. A. M. N. H. No. 10318. Twice natural size. ..... 60
Eleutherodactylus wightmanae. A. M. N. H. No. 10220. Twice natural size. 61
Eleutherodactylus richmondi. A. M. N. H. No. 10237. Twice natural size.... 63
Eleutherodactylus monensis. A. M. N. H. No. 24463 ....................... 65
Eleutherodactylus unicolor. Size of head (left), top of head (center), and inside
of mouth (right) of type....................... .................. 66
Eleutherodactylus unicolor. U. S. N. M. No. 26963, type. Twice natural size. 67
Digits of Hfemidactylus mabouia (left) and Sphaerodactylus macrolepis (right)
contrasted ....................... ............................. 68
Head of HIemidactylus mabouia ........................................ 70
Head and shoulders of Sphaerodactylus macrolepis. A. M. N. H. No. 13037 (A)
and No. 13697 (B), showing two common types of pattern. Two and a
half tim es natural size .......................... .................... 72
Heads of Porto Rican Anolis. Anolis cuvieri (left of top row), Anolis cristatellus
(center of top row), Anolis gundlachi (right of top row); Anolis evermanni
(left of middle row); Anolis stratulus (center of middle row), Anolis krugi (right
of middle row); Anolis pulchellus (left of bottom row), Anolis poncensis
(right of bottom row) ................... ........................ 76
Caudal crest of Anolis cuvieri (left), of A. gundlachi (center), and A. cristatellus
(rig h t) .... . . . ... .. . . . .. .. ... . . . . .. . .. . . .. . ... 78









CONTEXNTS OF 1 OLUI1E XV


Page
Dorsal scales of Porto Rican Anoles related to Anolis pulchellus. Left to
right: A. krugi, A. pulchellus, and A. poncensis ........................ 94
Head of Cyclura stejnegeri (type) ............ .......... ............. 103
Celestus pleii. A. 51. N. H. No. 13133. Natural size ...................... 106
Ameiva wetmorei. A. M. N. H. No. 13820. A. Head from above. B. Head
from side. C. Head from below. D. Arm from in front. E. Posterior
face of leg. F. Foot from above. G. Preanal scales. Three times natural
size ............................................ ........... 109
Ameiva wetmorei. A. 51. N. H. No. 13S21. Natural size................... 110
Ameiva alboguttata. A. M. N. H. No. 14003. Mona Island. Natural size... 116
Head of Amphisbaena caeca from above and from side.............. .... 118
Head of Amphisbaena bakeri from above and from side..................... 123
Head of Mabuya sloanii from above. A. M. N. H. No. 14007 (A) and A. M.
N. H. No. 14006 (B). To show variation in pattern. Twice natural size. 122
Head of AMabuya sloanii from side. A. M. N. H. No. 14007. Twice natural
size .......................................................... 122
Pattern of tail of Typhlops Platycephalus (left) contrasted with that of T.
rostellatus (right). A. M. N. H. Nos. 13336 and 13179. Natural size.. 126
Heads of Porto Rican Typhlops. Left to right (upper), T. platyecphalus; T.
rostellatus; (lower), T. monensis. (First two species from Stejneger; last
from Schm idt.) .................. ..................... .......... 127
Head of Epicrates inornatus from above (left), after Stejneger. Head of
Epicrates moneusis from above (right), after Schmidt ................ .. 131
Dromicus stahli, head from above and from below...................... .. 135
Color-pattern of Dromicus stahli ....................................... 136
Head of Dromicus exiguus from above and from the side.................... 137
Alsophis antillensis, head from above and from side ........................ 140
Color-pattern of Alsophis antillensis .................................... 140
Head of Alsophis portoricensis from above, showing scale-pits in dorsal scales.. 143
Color-pattern of Alsophis portoricensis. A. M. N. H. No. 8435. Twice natural
size ....................... ....................... ............ 144
Color-pattern of Alsophis variegatus. A. M. N. H. No. 13774............... 145
Carapace and plastron of Pseudemys stejncgeri. One-half natural size. ....... 148
Head of Pseudemys stejnegeri from below and from side, to show color-pattern.. 150
Branchiostoma caribaeum ............................................. 180
Asymmetron lucayanum. Johns Hopkins Univ. Studies. Biol. Lab. V. ...... 181
Ginglymostonma cirratum. From Zoologica, IX ........... . ......... 181
Galeocerdo tigrinus. From Zoologica, IX ................................. 182
Carcharhinus falciformis. From Zoologica, X ............................. 183
Carcharhinus limbatus, From Zoologica, X ............................... 183
Sphyrna zygaena. From Zoologica, IX . . . . . . . . . . . . . . . .... 184
Pristis pectinatus. Breder's Field Book of Marine Fishes (Putnam) .. ..... 185
Dasyatis americana. Breder's Field Book of Marine Fishes (Putnam) ........ 186
Dasyatis say. Breder's Field Book of Marine Fishes (Putnam).............. 187
Aetobatus narinari. From Zoologica, X .................................. 188
Anguilla rostrata. From Zoologica, IX . .......... ...................... 189
Leptocephalus conger. From Zoologica, X............. . ................... 190








vi S('IENTIFIC' SURVHEY OF PORTO RI('O

Page
M ayerina m ayeri .............................................. 190
M uraenesox savanna ........................................... . .. 191
A phthalmichthys caribbeus .............................................. 191
M yrophis longleii ......... ............................................. 192
Chilorhinus suensonii .......... ......................................... 192
Schagebranchus ophioneus ............................................... 193
Myrichthys oculatus. From Zoologica, X ............................. . 193
Myrichthys acuminatus. From Zoologica, X .............................. 194
Myrichthys keckii ...................................................... 194
Ophichthus ..., ................................................. 195
Gymnothorax moringa. From Zoologica, X............... .. .................. 196
Gymnothorax funebris. From Zoologica, X.. . ................ . 196
Gymnothorax albimnentis. . . . .. .......... .......... . . 197
Gymnothorax jordani ............................................. . 197
Echidna catenata................... ... ................... ........... 198
Tarpon atlanticus. From Zoologica, IX .............................. . 198
Elops saurus. From Zoologica, IX ...................................... 199
Albula vulpes. From Zoologica, IX ................................ . 200
Jenkinsia lamprotaenia. From Zoologica, X .......................... 201
Sardinella anchovia. From Zoologica, IX ............................. . 201
Harengula sordina. From Zoologica, X .................................. 202
HIarengula macrophthalma. From Zoologica, X ........................ . 203
Opisthonema oglinum. From Zoologica, IX ............................... 203
Anchovia perfasciata. From Zoologica, IX ............................. . 204
Anchovia brouwnii. From Zoologica, IX ............................. . 205
Anchovia choerostoma. From Zoologica, X ............................ . 205
Anchoria lyolepis. From Zoologica, X ........... ...................... 2C6
Cetengraulis edent lus. From Zoologica, X .......................... . .... 2C6
Trachinocephalus myoos. From Zoologica, IX ......................... . 207
Synodus intermedius. From Zoologica, X ............................ .. 207
Synodus foetens. From Zoologica, IX ................................ 208
Carassius auratus. .................. ................................... 209
Fundulus fonticola ...................................................... 210
Poecilia vivipara .................... .................................. 210
Tylosurus notatus ...................................................... 211
T ylostrus ardeola ........................... ............ .............. 212
Tylosurus raphidomna. From Zoologica, X ............................ . 212
Tylosurus acus. From Zoologica, IX ................................ 213
Hyporhamphus unifasciatus. From Zoologica, X ...................... . 214
Ilemiramphus brasiliensis. From Zoologica, IX ....................... . 214
Parexocoetus brachypterus. From Zoologica, IX ..................... . 2.. 15
Cypselurus bahiensis. From Zoologica, X ... . .. ............... . 215
Aulostomus maculatus. From Zoologica, X ........................... . 216
Fistularia tabacaria. From Zoologica, IX ............................ . 216
Syngnathus mackayi. From Zoologica, X ................................. 217
Syngnathus floridae........... ........................... ............ 217
Syngnathus elucens. From Zoologica, X .............................. 218
H ippichthys cayorum ............................................ ....... 218









(O7'TENXTI OF I OLUMI'E vii

Page
H ippichthys ensenadae .......................... ........ . . 219
Doryrhamphus sierra........................... .......... .......... 219
Hippocampus punctulatus. From Zoologica, X ............................ 220
Atherina stipes. From Zoologica, X.... .............................. 220
Atherina araea. From Zoologica, X . ....... ........................ 221
M ugil brasiliensis .................... ................. ............... 221
Mugil curema. From Zoologica, IX ............................... . 222
M ugil trichodon ..................... ............ ......... .......... 222
Agonostomus monticola. From Zoologica, X ......................... . 223
Sphyraena barracuda. From Zoologica, IX ........................... . 224
Sphyraena guachancho. From Zoologica, X ........................... 224
Sphyraena picudilla. From Zoologica, X .. . ........................ 225
Polynemus virginicus. From Zoologica, X ............................ . 225
MAfripristis jacobus. From Zoologica, X....... . ...................... 226
Holocentrus ascensionis. From Zoologica, X .. ........................... 226
Ilolocentrus vexillarius. From Zoologica, X .. ... .......... ......... 227
Upeneus maculatus. From Zoologica, X ..... .. ....................... 227
U peneus parous.............. ......... ...... .......... ............. 228
Upeneus martinicus. From Zoologica, X ................................. 228
Auxis hazard. From Zoologica, IX........ . . . ....................... 229
Scomberomorus maculatus. From Zoologica, IX ............. ....... .. 229
Scomberomorus regalis. From Zoologica, IX . . . . ..................... 230
Sconlberonmorous cavalla. From Zoologica, IX ........... .............. . 230
Trichiurus lepturus. From Zoologica, IX ................. ... ........ . 231
Oligoplites saurus. From Zoologica, IX ................ ..... ........... 232
Seriola falcata ......... .. .. ......... ........................ ........ 232
Decapterus punctatus. From Zoologica, IX ........................... 233
Trachurops crumenophthalmus. From Zoologica, IX ....................... 233
Caranx rubber. From Zoologica, X .......... .. ....................... 234
Caranx bartholomaei. From Zoologica, IX ......................... 235
Caranx hippos. From Zoologica, IX ....... ........................... 235
Caranx crysos. From Zoologica, IX ...... ............................. 236
Caranx latus. From Zoologica, X ....................................... 237
Vomer setapinnis. From Zoologica, IX ............................. 237
Vomer setapinnis cubensis. From Zoologica, X ........................ 238
Selene vomer. From Zoologica, IX ................................. . 239
Chloroscombrus chrysurus. From Zoologica, IX....... ..... ............ 239
Trachniotus glaucus. From Zoologica, X .... ............................ 240
Trachinotusfalcatus. From Zoologica, IX ............................ 241
Trachinotus carolinus. From Zoologica, IX .......................... . 241
Nomeus gronovii. From Zoologica, IX ............................... . 242
Peprilus parn. From Zoologica, IX .......... ........... ........... 243
Apogon sellicauda.............................. ..................... 244
Apogon conklitni ....................................................... 244
A pogonichthys alutus..................................... ............ 245
Apogonichthys stellatus. From Zoologica, X........ . ................ . 245
Centropomus undecimalis. From Zoologica, X ........................ .. 246
(entropomus parallelus........................ ....................... 246








viii <'IENTIFIC SU1R EY OF PORTO 11ICO

Page
Centropomus pectinatus. From Zoologica, X ........... ..... 247
Petrometopon cruentatus coronatus. From Zoologica, X ................ . 248
Cephalopholis fulrus ruber. From Zoologica, X .............. .............. 249
Cephalopholis fuitis punctatus. From Zoologica, X .................... ... 249
Epinephelus adscensionis. From Zoologica, IX ............................ 250
Epinepheluts striatus. From Zoologica, X ............................. 250
Epinephelus guttatus. From Zoologica, X ........................... 251
Epinephalus morio. From Zoologica, IX ............................. 251
Alphestes chloropterus. From Zoologica, X ............................ . 252
Mycteroperca bonaci. From Zoologica, IX .......................... . 252
M ycteroperca bowersi .................. ................................ 253
Hypoplectrus tnicolor. From Zoologica, X ............................ . 254
Prionodes baldwini .................. ..................... ............ 255
Dules dispilurus. From Zoologica, X ................................ . 256
Paranthias fu rcifer .................... .................. ............. 257
Rypticus coriaceus. From Zoologica, X .......... ......... ............ 258
Rypticus bistrispinus. From Zoologica, IX ........................... 258
Lobotes surinamensis. From Zoologica, IX .......................... . 259
Priacanthus arenatus. From Zoologica, IX ........................... . 260
Lutianus griseus. From Zoologica, IX ............................... 261
Lutianus jocu. From Zoologica, IX ................. .................... 262
Lttianus apodus. From Zoologica, IX ................................... 263
Lutianus aya. From Zoologica, X ....................................... 264
Lutianus analis. From Zoologica, IX ............... ........ ........... 265
Lutianus megalophthalmus ........... ...................... ............ 265
Lutianus synagris. From Zoologica, X ............................... 266
Ocyurus chrysurus. From Zoologica, X ................................. 267
Rhomboplites aurortbens. From Zoologica, X ............................. 268
A psilus dentatus......................................... ............ 268
Etelis oculatus ............................. ...... .................... 269
IIaemulon album. From Breder's Field Book of Marine Fishes (Putnam)..... 270
Haemulon macrostomum. From Zoologica, X......... . .............. 270
Haemulon bonariense. From Zoologica, X ............................ . 271
Haemulon parra .................... .................... .......... 272
Haemulon sciurus. From Zoologica, X ............................. . 273
Haemulon plumieri. From Zoologica, X................... .............. 274
Haemulon flavolineatum. From Zoologica, X .............................. 274
Bathystoma rimator. From Zoologica, X ............................. . 275
Bathystoma striatum. From Zoologica, X ............................. . 276
Anisotremus surinamensis ......................................... . 277
Anisotremus virginicus. From Zoologica, X ............................. 277
Pomadasys corvinaeformis. From Zoologica, X ........................ .... 278
Pomadasys crocro. From Zoologica, X ................................... 279
Calamus calamus. From Zoologica, X. .................................. 279
Calamus kendalli...................................... .............. 280
Calamus bajonado. From Zoologica, X ................................... 281
Calamus arctifrons. From Zoologica, X .................................. 281
Archosargus unimaculatus. From Zoologica, X ............................ 282









CONTENTT; OF 1 OLUME I ix

Page
Eucinostomvs pscudogula. From Zoologica, X ..... 283
Eucinostomus gula. From Zoologica, IX ...... . 283
Ulaema lefroyi. From Zoologica, X .......... 284
Xystaema cinereumn. From Zoologica, X ........ ... 284
Xyslaena havana ................. ............ ........ 285
Diapterus rhombeus. From Zoologica, X.................... 286
Diapterus olisthostonmus . . ................. .. 286
D iapterus brasilianus .. ... ................... 287
D iapterus plum ieri .......... .... ............ ........ 288
Kyphosus sectatrix. From Zoologica, IX .... .......................... 289
Cynoscion jamiaicensis. From Zoologica, X ............................. 289
Larimus brcviceps. From Zoologica, X ................................ 290
Odontoscion dentex. From Zoologica, X. ............................. .290
Bairdiella ronchus. From Zoologica, X . ... ........................ 291
Micropogon furnicri. From Zoologica, X ............................ 292
Eques punctatus. From Zoologica, X ..... ........................... 294
Eques lanccolatus. From Zoologica, X.. ... ........... 295
Pomaccutrus fascus. From Zoologica, X .. ... ..................... 304
Pomacentrus atrocyaneus .................. ...... .................. 305
Pomn acentrus analis ........... ......... ... .... ..................... 306
Pomacentrus leucostictus. From Zoologica. IX .. ... ...... ............ 306
Pom acentrus chrysus ............... ... .... .. ....... ........... 307
Abudcfduf saxatilis. From Zoologica, IX....... . . ................... 308
Abudefdaf analogus ..................... .. ............ .............. 309
Microspathodon chrysurus. From Zoologica, X ........................ 309
Microspathodon nimcatus. From Zoologica, X .................... ........ 310
M icrospathodon fowleri......................................... ........ 311
Lachnolaiimus maximus. From Zoologica, X .......................... 311
HIarpe rufa. From Zoologica, X ......................................... 312
Clepticus parrae. From Zoologica, X ............................... 313
Halichoeres garnoti. From Zoologica, X ............................. . 313
Halichoeres radiatus. From Zoologica, X. ........................... 314
Halichoeres bivitatus ................................... .... ........... 315
Halichocres kirschii ................. .. ............................. 316
Thalassoma nitidum. From Zoologica, X ................................. 316
Thalassoma bifasciatum. From Zoologica, X .......................... 317
Doratonotus megalepis. From Zoologica, X ........................... 317
Sparisoma radians. From Zoologica, X .................................. 318
Sparisonm a niphobles.. .................................... ............ 319
Sparisotna aurofrenatum. From Zoologica, X ......................... 319
Sparisoma abildgaardi. From Zoologica, X ........................... 320
Sparisoma chrysopterum. From Zoologica, X ............................. 321
Sparisoma lorito....................... .............................. 321
Sparisoma viride. From Zoologica, X .............................. . 322
Sparisoma flavescens. From Zoologica, IX ........................... . 322
Sparisoma rubripinne................... ............................. 323
Sparisoma brachiale. From Zoologica, X ............................. 323
Scarus taeniopterus. From Zoologica, X ............................ . 324








x NCIESTIFIC SNiR1 El OF I'ORTO RICO

Page
Scarus punctulatus. From Zoologica, X .......................... .... 325
S cars vetula ............................................ ........... 325
Scarus croicensis. From Zoologica, X .................................... 326
Scarus coeruleus............... ........ ................. ............. 327
Pseudoscarus guacamaia. From Zoologica, X ............................. 327
Chaetodipterus faber. From Zoologica, IX ............................ . 328
Chaetodon ocellatus. From Zoologica, IX ............................. 329
Chaetodon striatus. From Zoologica, X ............................... 330
Chaetodon capistratus. From Zoologica, IX ........................... . 331
Pomacanthus arcuatus. From Zoologica, IX .......................... 332
Holacanthus tricolor. From Zoologica, X ............................. 332
Angelichthys ciliaris. From Zoologica, X ............................ 333
Teuthis caeruleus. From Zoologica, IX ............................. 334
Teuthis hepatus. From Zoologica, IX .................................... 334
Teuthis bahianus. From Zoologica, IX ................................... 335
Balistes vetula. From Zoologica, IX ..................... ............ 336
M elichthys piceus ........................................ ........... 337
X anthichthys ringens.................... ................. ............ 338
Cantherines pullus. From Zoologica, X .............................. 339
Cantherines am phioxys. .............. ..................... .......... 339
Monacanthus ciliatus. From Zoologica, X ............................ . 340
Monacanthus tucker. From Zoologica, X .................... ....... . 341
Monacanthus hispidus. From Zoologica, IX .......................... 341
Alutera script. From Zoologica, X ............................. ... . 342
Lactophrys triqueter. From Zoologica, IX..... . .............. . 343
Lactophrys trigonus. From Zoologica, IX ........................... . 344
Lactophrys bicandulis. From Zoologica, N .... .......................... 345
Lactophrys tricornis. From Zoologica, IN .................. .............. 345
Lagocephalus laevigatus. From Zoologica, IX ......................... 346
Tetraodon spengleri. From Zoologica, IX .............................. 347
Tetraodon min rmoratus. From Zoologica, X ........................... 347
Tetraodon testudineus. From Zoologica, IX ........................... 348
Canthigaster rostratus. From Zoologica, X ........ ...................... 349
Diodon hystrix. From Zoologica, IX ............................... . 349
Diodon holacanthus. From Zoologica, X ............................. . 350
Chilomycterus antennatus. From Zoologica, X. ......................... . 351
Scorpaena brasiliensis. From Zoologica, X.......... ................ . 351
Scorpaena albifim bria.................................... ............ 352
Scorpaena bergii.............. ...... .. ................ ............. 353
Scorpaena plumieri. From Zoologica, IX ............................. . 353
Scorpaena grondicornis. From Zoologica, IX ......................... . 354
Pontinus beanorum........... ..... ................................... 354
Pontinus macrolepis ................ .. ............................. 355
Prionotus punctatus. From Zoologica, X ............................. . 356
P eristedion gracile............................ ......... .. ....... ...... 357
Ccphalacanthus volitans. From Zoologica, IX ......................... . 357
Collionymius calliurus ................................... ............. 359
Gobiomorus dormitor. From Zoologica, X ........................... ..... 359









t'O)NTENT' OF VIOLI'1E X xi

Page
Dormitator maculatus. From Zoologica, X ................................ 360
Guavina guavina ...................................................... 361
Eleotris pisonis. Breder's Field Book of Marine Fishes (Putnam)........... 361
Sicydium antillarum ......... .................................... 362
Sicydium caguitae ....... ............................................ 362
Sicydiumw n plnmnieri...................................................... 363
Bathygobius soporator. From Zoologica. X ........................... . 363
Gobius translucens .......... ........................................ 364
Gobius glaucofraenum .................................... .......... .. 365
Gobius bolcosom a........................................... .......... 365
Gobius lyrics ................ .................................. 366
G obius bayam onensis .................... ............................. ... 366
Gobius oceanicus. From Zoologica. X .......... ......................... 367
Chonophorus taiasica. From Zoologica, X ............................ . 367
Bollmannia boqueronensis ............................................... 368
M icrogobius m eeki................ ....................... ............ 368
Gobiosoma multifasciatum ............................................... 369
Gobioides broussonnetii. ........... .................. ............. ..... 369
Echeneis naucrates. From Zoologica, IX ................................. 370
Malacanthus plumieri. From Zoologica, X ........................... . 371
Caoulolatilus cyanops ......... ......... ........................... ..... 372
Dactyloscopus tridigitats. ................................................ 372
Gobiesox tudes......... .... .. .......................................... 373
Gobiesox cerasinus......................... ......................... 373
Gillias jordani. From Zoologica, X ...................................... 374
Brannerella culebrac. From Zoologica, X ............................. 375
A cteis m oorei. From Zoologica, X ....................................... 375
M alacoctenus ptertoricensis .............................................. 376
M alacoctenus delalandi........................................... . 376
Labrisomus nuchipinnis. From Zoologica, X ......................... . 377
Auchenopterus albicandus ............................. .... ............. 378
Auchenopterus fajardo. From Zoologica, X ........................... 378
Auchenopterus rubescens ............................. ............. 379
Tekla cingulata ..................... ................................... 379
Tekla fasciata......................... .................. .......... . 380
A uchcnistius stahli. From Zoologica, X .............................. 381
Bupiscartes macclurei.......... ....................................... 381
Blennius cristatu s. Breder's Field Book of Marine Fishes (Putnam) ......... 382
Salarichthys textilis ........ ................ ............................ 382
Coralliozetus cardonae .............. ................................. 3S3
Emblemaria pandionis ............................................ ..... 383
Fierasfer berm udensis .................... .. .............. ............. 384
Platophrys occllatus. From Zoologica, IX ............................ . 385
I'latophrys lunatus. From Zoologica, X .................................. 386
Syacinm niicrurmn. From Zoologica, X .. . . ..... .................. 387
Citharichthys unicornis................ ................... ........... 387
Citharichthys spilopterus. From Zoologica, X ..................... ...... 388
Citharichthys arenaceus ............... .................................. 389








,CIENTIFlC ,s'ti'IEY OF P'ORTO RICO


Page
E trop us crossotus ...... ........................... . .. . . .. 389
A chirus inscriptus ......................... ........ .. ...... 390
Achirus lineatus. From Zoologica, X ................. .................. 390
Symphurus plagusia. From Zoologica, X ............................. . 391
H istrio gibbus. From Zoologica, X ...................................... 392
Antennarius inops. From Zoologica, X ................................. 393
Antenmarius scuber ............. ...................................... 394
Antennario s nuttingii ........ .. ...................................... 394
A ntennarius m ultiocellatus ... .......................................... 395
Chaunax pictus....................... ... ............................ 395
Ogcocephalus vespertilio. From Zoologica, IX ........................... 396
Halieutichthys aculeatus. From Zoologica, X ........................... 397
Halieutichthys smithii ............... ............................... . 398
Internal Anatomy of Ascidian ........... ........... ................... 407
Comparative diagrams of the tadpoles or larval stages of an ascidian (upper
figure), and of a frog (lower figure), to show their correspondence in many
points of structure, especially in having gill clefts in the walls of the pharynx
or throat and a rodlike notochord corresponding to the backbone in the
higher animals, above which is the main part of the nervous system, cor-
responding to the brain and the spinal cord .......................... ... 409
Polyclinun constellatum Savigny, 1816. Zooid, X30, and part of the surface
of a colony, showing the arrangement of zooids and common cloaca] canals,
X 4 .6 .. .............. .... .......... ..... .. ......... ............ 423
Aplidium lobatum Savigny, 1816. Zooid, X32 ........................ ... 425
Aplidium (Amaroucium) bermudae (Van Name). Three colonies, slightly
enlarged ....... .... ....................... ....... ............... 426
Aplidium (Amaroucium) bermudae (Van Name), 1902. Zooid, X25.......... 427
Trididemnum savignii (Herdman), 1886. A colony, natural size ............. 429
Trididemnumnt ..-u,.., (Herdman), 1886. Zooid, X40, and groups of spicules,
X 460 .. ............................ ....... ....................... 430
Trididemnum solidum (Van Name), 1902. A colony attached to dead coral,
natural size ................... ... .......... .... ................ 431
Trididemnum solidum (Van Name), 1902. Zooid, X60, and spicules from two
colonies, X 460 ......................... .......... ................. 432
Trididemnum orbiculatum (Van Name), 1902. A living colony magnified
nearly X3 ...................................................... 433
Trididemnum orbiculatum (Van Name), 1902. Zooid, X40, spicules X460;
part of the surface of a colony to show the distribution of spicules in the
superficial layer of the text X 11 ....................... ............... 434
Didemnwnu candidumn Savigny, 1816. A colony, natural size................. 435
Didemnun candidum Savigny, 1816. Zooid, X40, and groups of spicules show-
ing their variation in different colonies, X460 ........................... 437
Didemnumn (Polysyncraton) amethysteum (Van Name), 1902. A colony attached
to a sponge, natural size.............................. .............. 439
Didemnum (Polysyncraton) amethysteum (Van Name), 1902. Zooid, X40.
Spicules (all from same colony), X460, and part of the surface of a colony
showing distribution of spicules in superficial layer of test, Xll ............ 440








COXTEXTN' OF VOLUME -


Page
Diplosoma miacdonaldi Herdman, 1886. A colony, natural size ........... 441
Diplosoma macdonaldi Herdman, 1886. Zooid, X48 ....................... 441
Lissoclinum fragile (Van Name), 1902. Zooid, the branchial sac well extended,
X36, and spicules (all from the same colony) X460 ...................... 442
Echinoclinum verrilli (Van Name), 1902. A colony, X2; zooid, X36; group of
spicules, X230 .................................................... 444
Polycitor (Eudistonma) olivaceus (Van Name), 1902. Two colonies, natural size. 445
Polycitor (ERdisloma) olivaceus (Van Name), 1902. Zooid, X32. ............ 446
Polycitor (Eudistoma) clarus (Van Name), 1902. Zooid containing four de-
veloping em bryos or larvae, X 35...................................... 419
Clavelina oblonga Herdman, 1880. Two colonies, showing zooids almost com-
pletely separated (left figure) and one section of a completely consolidated
colony (right figure), both natural size ............................. . 451
Clavelina oblonga Herdman, 1880. A small colony with separate zooids, X 1.8,
and three lobes of a large colony with partially united zooids, X 1.8 ....... 451
Clavelina oblonga Hermand, 1880. Zooid branchiall sac contracted: embryos
and larvae in peribranchial cavity), X 10.5.............................. 452
Cystodytes dellechiaiae (Della Valle), 1877. Two entire colonies and a section
of another colony, natural size..................... ................ 453
Cystodytes dellechiaiae (Della Valle), 1877. Zooid, X32, and a group of spicules,
X 52 . ....... .. .... .... . .. . ..... .............. ... ........... 453
Distaplia bermvdensis (Van Name), 1902. A colony, natural size............... 454
Distaplia bermudensis (Van Name), 1902. Zooid with incubatory pouch con-
taining three embryos or larvae, X30 .................................. 455
Distaplia bursata (Van Name), 1921. Three colonies, X I.3, and zooid X55. 457
Rhopalaea abdominalis (Sluiter), 1898. On the left, the outline of the entire
individual, including the test, slightly enlarged. In the center, the individual
removed from test, showing muscle bands on mantle, X5.6; also a piece of
the branchial sac, X5.6. In the right, the upper part of the median dorsal
vessel, showing the dorsal tubercle and five of the dorsal langucts, X56. . ..458
Perophora viridis Verril, 1871. Zooid, X27 ........................... ... 460
Ecteinascidia turbinata Herdman, 1880. Zooid, X<7.2.................... 462
Ascidia nigra (Savigny), 1816. The left side of the body slightly reduced; the
dorsal tubercles of two individuals and part of the branchial sac, X32 ...... 463
Ascidia hygomiana (Traustedt), 1882. Natural size ........................ 464
Ascidia hygomiana (Traustedt), 1882. The left side of the body, X3, dorsal
tubercle, X10, and part of branchial sac, X36........................... 465
Ascidia curvata (Traustedt), 1882. The left side of the body, X2.2, dorsal
tubercle, X16, Fig. 68, and part of the branchial sac, X42 ................ 467
Ascidia sydneiensis Stimpson, 1885. The dorsal tubercle, X12; the left side
of the body, X1.6; the right side of the body showing the arrangement of
the muscles in the mantle, X 1.6; and part of the branchial sac, X42....... 468
Ascidia corelloides (Van Name), 1924. The left and right sides of the body,
X3.5 ............................................. ............ 469
Ascidia corelloides (Van Name), 1924. Part of the branchial sac, X25, and
part of the circle of tentacles with dorsal tubercle and anterior end of the
branchial sac, X IS ................................................... 470


xiii







('IENT'IFIC SU'RHi'EY OF PORTO RICO


Page
Ascidiella styeloides (Traustedt), 1882. The left side of the body, X4, and
part of the branchial sac, enlarged (outlines of Traustedt's figure).......... 471
Rhodosoma turcicum (Savigny), 1816. The left and right sides of the body, X2,
and part of the branchial sac, X38..................................... 472
Corella minute (Traustedt), 1882. The left and right side of the body, X2.5,
and part of the branchial sac, X25..................................... 473
Botryllus plans (Van Name), 1902. Zooid seen from left side, X36 ......... 476
Botryllus schlosseri (Pallas), 1766. A colony, natural size ................. 478
Botryllus schlosseri (Pallas), 1766. Zooid, X36 ....................... . 478
Botrylloides nigrurn Herdman, 1886. Zooid, X40............. ........... 480
Outlines of the stomach of (a) Botryllns plans (Van Name), 1902, and (b)
Botrylloides nigrum Herdman, 1886.................................. 481
Symplegma viride Herdman, 1886. A colony attached to rock. Natural size.. 483
Symplegma viride Herdman, 1886. Zooid (individual with developing eggs),
X35 ......... ................. ............................... 483
Polyandrocarpa (Eusynstyela) lincta (Van Name), 1902. Part of the branchial
sac, X45, and left (upper figure) and right (lower figure) sides of zooid, X9.. 485
Polycarpa obtecta Traustedt, 1883. Two specimens, natural size............. 487
Polycarpa obtecta Traustedt, 1883. The left and right sides of the body, natural
size . .. . ... . . . .. .. .. . . . .. .. . .. .. . .. .. . . . .. .... . 4 87
Polycarpa spongiabilis Traustedt, 1883. Natural size ...................... 488
Styela partita (Stimpson), 1852. Specimens natural size. ................... 490
Styela partita (Stimpson), 1852. The left and right sides of the body, X2.
The dorsal tubercle, X12. A gonad of a small individual, X20, and part
of the branchial sac, X 14.................. ........... .............. 491
,Styela plicala (Lesueur), 1823. Four specimens, natural size. ............... 493
Styela plicata (Lesueur), 1823. Upper figures: left and right sides of body,
slightly enlarged, and terminal part of a gonad of an individual having the
gonads compact and the testes of simple form, X16. Lower figures: dorsal
tubercle, X6, and terminal part of the gonad of an individual with highly
developed branching testes, X 12 ....................................... 494
Pyura vittata (Stimpson), 1852. The body removed from the test, X1.5..... 496
Pyura vittata (Stimpson), 1852. The left and right sides of two individuals.
The large one is only slightly enlarged; the small one is 1.6 times the natural
size ............................................................ 496
Pyura vittata (Stimpson), 1852. Upper figures: dorsal tubercles and tentacles
of different individuals to show variation, X12 to 20 times. Lower figures:
genital sacs from gonads of two individuals, one sac fully, the other partly,
filled by the reproductive glands, X14; also (in center) part of the liver,
X 18 ........................................................... 497
Pyura momus form pallida (Heller), 1878. The left and right sides of the body,
natural size ................... ...................................... 498
Pyura momus form pallida (Heller), 1878. Upper figures: spicules. On the left,
part of a spicule, X420; on the right, spicules from the test (small group) and
from the vessels of the branchial sac (large group), X35. Lower Xfigures:
anatomy. On the left, dorsal tubercle, X8, and tentacle, X25; on the right,
part of a gonad, X8, and part of liver, X20....................... .. 499








CONTENT'I OF 1 OLUfME


Page
IMicrocosmus claudicans exasperatus (Heller), 1878. Two specimens, natural
size. The right-hand one has three species of compound ascidians growing
on it. At a, Clavelina oblonga; below b, Dislaplia bermudensis; opposite c,
Didem num candidum ............................. .................... 501
Microcosmus claudicans exasperatus (Heller), 1878. The body removed from the
test is slightly enlarged: the right-hand specimen is transversely sectioned to
show the folds of the branchial sac................................... 501
Microcosmus claudicans exasperatus (Heller), 1878. The left and right sides
of the body, X 1.2......................... ................... .. 501
Microcosmus claudicans exasperatus (Heller), 1878. Upper figures: intestinal
loop and gonad of left side, seen from side next to the branchial sac, X6, and
tentacles, X15. Lower figures: dorsal tubercles of two individuals, X7, and
part of liver, X10, and minute spines from the lining of the distal part of the
branchial tube, X 280............. .......... ........................... 502
Microcosmus helleri Herdman, 1881. Four specimens, natural size. .......... 504
Microcosmus helleri Herdman, 1881. The left and right sides of the body, X1. 5. 504
Molgula occidentalis Traustedt, 1883. Natural size ..................... ... 506
Molgula occidentalis Traustedt, 1883. The left and right sides of the body, X 1.5 506
Molgula occidentalis Traustedt, 1883. On the left, large tentacle, X9, the
dorsal tubercle, X12, part of the liver, X6, and the closed end of the left
gonad, X15. On the right, part of the branchial sac, X12 ....... ....... .507

PLATES

1. The Arid Section of Porto Rico. View east of Ponce.
The Arid Extreme in Porto Rico. View near Guanica.
II. Open Place in Sierra Palm Forest, El Yunque. A colony of Begonia
decandra occupies the center; the trunks of the palms are covered with
mosses, juvenile plants of Maregravia, and a few large Bromeliaceae.
(From Plant Ecology of Porto Rico, by H. A. Gleason and Mel T. Cook.)
III. Forests of the Upper Slopes of El Yunque. Altitude about 1050 m. The
distribution of sierra palms in strips and patches follows the contour of
the land. (From Plant Ecology of Porto Rico by H. A. Gleason and
Mel T. Cook.)
IV. The Porto Rican Boa, Epicrates inornatus Reinhardt. Luquillo Forest.
Photographed by F. H. Winslow; photograph supplied through the
courtesy of B. A. Wall.
V. Clavelina oblonga Herdman, 1880. A colony attached to a gorgonian,
X about 1.3 (From Trans. Conn. Acad. Sci., Vol. XI, Pl. LX).
VI. Ecteinascidia turbinata Herdman, 1880. A colony attached to a mangrove
root. Nearly natural size.
VII. Ascidia nigra (Savigny), 1816. Three individuals, natural size (upper
figure). Polytcitor hepaticus Van Name, 1921, natural size (lower figure).
VIII. Botryllus plans (Van Name), 1902. Two differently colored living
colonies growing on the same rock; X more than 2.5. (From Trans.
Conn. Acad. Sci., Vol. XI, Pl. LXI.)





/







AMPHIBIANS AND LAND REPTILES OF
PORTO RICO

With a List of Those Reported from the Virgin Islands

BY KA III PATTERSON SCHMIDT

CONTENTS
Page
Introduction ........................ ..................... ............ 3
Itinerary and collections m ade........... .......... .. ........... 4
Other m material exam ined............................. .............. 4
Plan of work .............................. .......... ............. 5
Acknowledgments ..................... ............................ 6
Porto Rican herpetology since 1904 ................................ 6
Lists of the amphibians and land reptiles of Porto Rico and the adjacent
islands .......... ......................... ......... ............ 7
Habitat associations and faunal subdivisions .......................... 9
Origin and relations of the Porto Rican herpetological fauna ............ 12
Systematic account of the species ........................................ 30
Class Amphibia ............................ ..................... 30
Order Salientia ........................ .......... ........... 30
Fam ily Bufonidae.................... ....... ............... 30
Key to the genera of Porto Rican frogs and toads. ......... 30
B ufo Laurenti ............... ....... .... ............ 31
Key to the Porto Rican species of true toads .............. 31
Bufo lemur (Cope) ................ ....... ............ 31
Bufo marinus (Linnd) .................................. 34
Leptodactylus Fitzinger. ............ .................. 37
Leptodactylus albilabris (Giinther) ...................... 37
Eleutherodactylus Fitzinger .......................... 43
Eleutherodactylus portoricensis Schmidt.................... 44
Elcutherodactylus gryllus Schmidt ...................... 51
Elevtherodactylus locustus Schmidt........................ 53
Elcutherodactylus cram ploni Schmidt...... ............... 55
Eleutherodactylus antillensis (Reinhardt & Luetken) ........ 56
Eleutherodactylus brittoni Schmidt ...................... 59
Eleutherodactylus wightmanae Schmidt .................... 60
Eleutherodactylus richmondi Stejneger..................... 62
Eleutherodactylus monensis (Meerwarth) ................... 65
Elevthcrodactylus unicolor Stejneger ................... 66
C lass R eptilia ........................ ........ ................... 6S
Order Squam ata ................... .......................... 6S
Suborder Sauria ............................... ......... 68
Key to the genern of Porto Rican lizards ............. 68
Gckkonidae ................... ......... ............ 69
Ilem idactylis O ken ......... ...... .. ........... 69







2 SCIENTIFIC SURVEY OF PORTO RICO

Page
Hemidactylus mabouia (Moreau de Jonnes) ............ 69
Sphaerodactylus W agler ............................. 70
Sphaerodactylus macrolepis Giinther .................. 70
Iguanidae................... ........... ........... 74
A nolis D audin..................................... 74
Key to the species of Anolis recorded from Porto Rico. 74
Color key to Porto Rican anoles ..................... 75
Anolis cuvieri M errem ............................. 76
Anolls cristatellus Dumeril and Bibron................ 80
Anolis gundlachi Peters ........................... 85
Anolis stratulus Cope ............................. 87
Anolis evermanni Stejneger.......................... 90
Anolis pulchellus Dum6ril and Bibron ................ 92
A nolis krugi Peters.................. .............. 96
Anolis poncensis Stejneger.......................... 99
Cyclura H arlan...................... ............ 101
Cyclura stejnegeri Barbour and Noble ................ 102
Celestus G ray....................... ............. 105
Celestus pleii Dum6ril and Bibron.................... 105
T eidae ................... ........... .... ........... 108
Ameiva M eyer...................... ............. 108
Synopsis of the Porto Rican Ameivas ................. 108
Ameiva wetmorei Stejneger ...................... . 108
Ameiva exsul Cope................................. 112
Ameiva alboguttata Boulenger ........................ 115
Am phisbaenidae ............ ........... ............. 117
Amphisbaena Linnd ............................. 117
Amphisbaena caeca Cuvier ........................ 117
Amphisbaena baker Stejneger ....................... 119
Scincidae ....................... .. ................ 121
M abuya Fitzinger ................................. 121
M abuya sloanii (Daudin) .......................... 121
Suborder Serpentes......................................... 124
Synopsis of the genera of Porto Rican snakes.......... 124
Typhlopidae.......................... ............... 124
Typhlops Oppel .............. ..................... 124
Key to the Porto Rican species of Typhlops. .......... 124
Typhlops platycephalus Dum6ril and Bibron. .......... 125
Typhlops rostellatus Stejneger... . ........... . ... . 128
Typhlops monensis Schmidt ......................... 129
B oidae............................... .............. 130
Epicrates W agler................................... 130
Synopsis of the Porto Rican and Mona Island Epicrates.. 130
Epicrates inornatus (Reinhardt)...................... 130
Epicrates monensis Zenneck ......................... 132
C olubridae ............. ... .......... .............. 133
Dromicus Bibron.................. ................ 133
Dromicus stahli (Stejneger) .......................... 134
Dromicus exiguus Cope ............................ 137
Alsophis Fitzinger.................. ............... 139







SCHMIDT, AMPHIBIANS OF PORTO HICO


Page
Alsophis antillensis (Schlegel) ....................... 139
Alsophis portoricensis (Reinhardt and Luetken). ....... 142
Alsophis variegatus (Schmidt)........... ........... .. 144
Order Testudinata.............. .. .................... 147
Emydidae ............ ............................. 147
Pseudemys Gray ................................. 147
Pseudemys stejnegeri, sp. nov .................... 147
A hand-list of the amphibians and reptiles of the Virgin Islands.............. 150
Distributional list of the amphibians and reptiles of the Virgin Islands. . 151
Notes on herpetology of the Virgin Islands .................. ......... 151
Artificial keys to the species ................................ . 153
Bibliography............... .................................. 155


INTRODUCTION

The new account of the amphibians and reptiles of Porto Rico and its
dependent islands here attempted, while based on the fundamental work
of Stejneger (1904), has an independent source in the collections made
in the course of the Scientific Survey of Porto Rico and the Virgin
Islands undertaken by the New York Academy of Sciences, in coopera-
tion with the Porto Rican government and The American Museum of
Natural History. These collections consist of a total of 1435 speci-
mens, representing 33 of the 42 forms in the area covered by this report.
In the course of investigations on other groups of animals, 103 speci-
mens of amphibians and reptiles were collected by F. E. Lutz, J. T.
Nichols, R. W. Miner, H. E. Anthony and T. H. Jones previous to 1919.
It was my good fortune to conduct the first specifically herpetological
field-work for this Survey in the summer of 1919, and for this opportu-
nity I am indebted primarily to Miss Mary C. Dickerson, then Curator
of Herpetology at The American Museum of Natural History, and to
Dr. Henry C. Crampton, Dr. N. L. Britton and Dr. Ralph W. Tower, of
the Porto Rico Committee of the New York Academy of Sciences. In
carrying out this field-work, Mrs. Schmidt and I spent the period from
August 3 to October 8 in investigations on Porto Rico and in mak-
ing visits to the adjacent islands-Mona, Vieques and Culebra. Our
collections amounted to 1253 specimens.
These collections were considerably enriched in 1926, by Messrs. II. E.
Anthony and G. G. Goodwin, who collected 74 specimens on Mona
Island, and 5 on Caja. de Muertos, southeast of Ponce. This material
includes a fine series of Eleutherodactylus monensis, which was wanting
in my Mona Island collection, and the first herpetological specimens
from Muertos Island.






SCIENTIFIC SURVEY OF PORTO RICO


ITINERARY AND COLLECTIONS MADE

Mrs. Schmidt and I arrived at San Juan on August 3rd, 1919. The
first week of our stay was spent at Santurce, which provided a con-
venient base for the necessary official visits in San lJuan. There we
were able also to do some introductory collecting, with our interest
stimulated by a tree frog new to Porto Rico that was abundant in the
hotel grounds. lRio Piedras and Cataiio were also visited from San-
turce. The next two weeks (August 11-24) we sojourned at Aibonito,
at an altitude of about 2000 feet, in the heart of the coffee-belt. Daily
excursions were made in this vicinity. Four days were spent at Coanmo
Springs, a truly delightful collecting locality, affording numerous spe-
cies new to our collections.
On( August 29 we returned to Santurce for a fresh start. With Mona
Island as objective I went alone to Mayagiiez (September 3). There
I learned that it would lie impossible to sail for Mona until September G.
This delay enabled me to make a productive trip to Maricao. Thle week
of September 7-13 was spent in the trip to Mona Island, on which I
was accompanied by E. M. Bruner, Forester of Porto Rico.
Returning to Santuree, where specimens were meanwhile aceumulat-
ing, thanks to the interest of B. A. Wall, of Bayanmon, we packed and
stored the collection. On September 18 we left by) rail for Ensenada,
where we enjoyed the hospitality of Superintendent Boyd, of the
Giuanica Central, at "Canary Cottage."
Again returning to Santurce (September 28) I set forth on a three-
days trip to El Yunque, where I camped in the Forester's Cabin at 1200
feet altitude, climbing to the peak on September 30. This was followed
by a brief trip to Vicques and Culebra islands by means of a sloop
chartered at Fajardo. On October 8 we sailed from San Juan for New
York.
OTHI:ni MAITEIZIAL EXAMINED '

Besides the material collected by the Survey of Porto Rico and the
Virgin Islands, all of which is deposited in The American Museum of
Natural History, I have had the privilege of examining, thanks to the
courtesy of Dr. W. C. F. McClure, the Porto Rican collection preserved
at Princeton University.
Dr. Stuart T. Danforth, of thie University of Porto Rico, at Maya-
giiez. kindly sent me both his personal collections and those of the Uni-
versity for examination in connectioii with this report. ,.',








SCHMIDT, A.1PlIB!IANS OF PORTO. RICO


PLAN OF WORK

"The Herpetology of Por.fo Rico" by Dr. Leonhard Stejneger (1904)
is a work of exceptional merit. It remains a model for the exact and
complete description of an insular fauna, and sets a high standard for
systematic zoology in general. It is a pleasure to record here the use-
fulness of this volume. A copy accompanied me to Porto h'ico in 1919
and proved most serviceable as a field manual, making possible thl
identification of most of the species and thus facilitating all phases of
field study.
It was my first plan to prepare merely a supplement to Dr. Stejneger's
report, embodying only the additions to our knowledge of the Porto
Rican herpetological fauna since 1904. After a review of the necessary
additions, in conference with Dr. H. ('. C'rampton, it was decided, how-
ever, to enlarge the scope of the work and present a renewed "complete
account" both for the sake of increased usefulness to future students and
to bring it into better accord with the similarly complete reports of
other contributors to the Survey. The existence of Stejueger's report
has greatly simplified the preparation of the present one. In the
case of the numerous species whose definition has required no change.
I have followed Stejneger's descriptions closely or quoted them ver-
batim, and I have availed myself of a large number of his text figures.
especially for the illustration of key characters. The figures drawn for
the present paper are designed to present the habits of a number of
species, and thus supplement Stejneger's otherwise complete illustra-
tion of the fauna. These figures are the work of Mrs. E. Beuten-
miuller, whose drawings have embellished so many herpetological papers.
The half-tone figure of Eleutherodaritylus muicolor was supplied through
the courtesy of Dr. Stejneger.
I have adopted a conservative position on one phase of nomenclature.
Excellent arguments might be advanced for treating several of the Porto
Rican forms as subspecies rather than as full species. Such a nomen-
clature would reflect more information as to the actual relations of the
forms concerned than binomial treatment. The species of Typhlops
allied to jamtalcensis, the fresh water turtle, and the Mima Island
Aime'ra and Cyclura are eases in point. It is very difficult, however, to
draw a line between insular subspecies and insular species, and our
knowledge of many forms is manifestly imperfect. Any attempt at a
trinomial arrangement of Eleutlieroducltylus is obviously impossible. I
have accordingly left the matter for future consideration, preferably in
connection with a new list of the West Indian fauna as a whole.








SCIENTIFIC SURVEY OF PORTO RICO


So much work still remains to be done on the herpetological fauna
of Porto Rico by some resident naturalist, especially with reference to
the discrimination of the small tree frogs and their life .histories, that
the present account of the fauna is hardly more likely to be "final" than
was that of Dr. Stejneger more than twenty years previously.

ACKNOWLEDGE ENTS
It is a pleasure to acknowledge the cordial furtherance of the present
work by the members of the Porto Rico Committee of the New York
Academy of Sciences, by my various sometime colleagues of The Ameri-
can Museum of Natural History who took part in the Survey of Porto
Rico and the Virgin Islands, and by nearly everyone with whom we
came in contact in the course of the herpetological field-work.
We were especially indebted, when in Porto Rico, to MIr. and Mrs.
B. A. Wall, of Bayamon ; to Mr. E. M. Bruner, Forester of Porto Rico;
to Mr. Marc Lejeune, of Mayagiiez, to whom I owe the visit to Mona
Island: and to Colonel George A. Shanton, Chief of the Insular Police.
In the course of the preparation of the report I have had the most
cordial aid from various herpetologists. I have applied for information.
for specimens or for advice to Dr. Leonhard Stejneger and Miss Doris
Cochran, of the United States National Museum: to Dr. Thomas Bar-
hour and Mr. Arthur Loveridge, of the Museum of Comparative Zool-
ogy; to Dr. G. K. Noble and Mr. Clifford H. Pope, of The American
Museum of Natural History; to Dr. Emmett Reid Dunn, of Smith Col-
lege: to Dr. Stuart T. Danforth, of the University of Porto Rico; and
to Mr. H. W1. Parker. of the British Museum (Natural History).
The friendly criticism and interest of Mr. Herbert F. Schwarz, now
editor of the reports of the Survey of Porto Rico, have improved the pres-
ent paper at innumerable points, both in minor details and in more im-
portant matters. My thanks (and still more those of the reader) are
due to him for great patience with a difficult manuscript.

POnTO RICAN HERPETOLOGY SINCE 1904
Stejneger presents an excellent historical review of the growth of our
knowledge of the amphibians and reptiles of Porto Rico (1904, pp. 553-
5;'9). The small but interesting collection secured by W. W. Brown,
Jr., on Mona Island in February. 1892, has since come to light and was
reported on by myself (1926).
Subsequent to the collections made for the United States National
Museum in 1899-1901, no mention of Porto Rican herpetology appeared








SCHMIDT, AMPHIBIANS OF PORTO RICO


until 1913, when Stejneger described the unusually interesting and ex-
tremely distinct A meira icetmorei from Rio Loco, near Guanica. The
type was collected by Dr. Alexander Wetmore in the course of his in-
vestigations of the Porto Rican bird fauna.
The collections made by Mr. Charles F. Silvester, while on the staff
of the expedition of the Carnegie Institution to Porto Rico in 1915, were
reported upon by Fowler in 1918. Fowler figures Ameiva wetniorei
and discusses variation in other species.
The discovery of bones referable to an extinct species of Cyclura in a
cave near Ciales by Dr. Glover M. Allen and James Lee Peters, in 1917,
filled an important gap in the distribution of this typically Greater
Antillean genus. The species was described by Barbour (1919), the
type being the extremities of a left humerus, with numerous additional
limb-bones, jaws and vertebrae. Similar material was collected for the
Scientific Survey of Porto Rico and the Virgin Islands by II. E. An-
thony in 1916.
The herpetological collecting of the various workers who have taken
part in the Scientific Survey of Porto Rico and the Virgin Islands has
been described above.
Dr. E. Greywood Smyth, Entomologist for the Porto Rican Agricul-
tural Experiment Station at Rio Piedras, has paid some attention to the
amphibians and reptiles and in 1920 published a brief account of the
food habits of the Anoles.
The food habits of Porto Rican lizards were subsequently analyzed in
some detail by George N. Wolcott, in a paper published in 1924 in the
Journal of the Department of Agriculture of Porto Rico.
A small collection made in the course of ornithological investigations
in 1924-1925 was reported upon by Stuart T. Danforth (1925 and
1926). This material was subsequently purchased by the Field Mu-
seum of Natural History. AMr. Danforth has also collected on Desecheo
Island, adding Ameiva exsul to the list from that island in 1926.

LISTS OF THE AMPHIBIANS AND LAND REPTILES OF PORTO Ri'O AND
THE ADJACENT ISLANDS

I. PORTO RICO
1. Bufo lemur 5. Eleutlherodaclylus gryllus
2. Bufo mna-rinus* 6. Eleutlerodactylus locustus
3. Leptodactylus albilabris 7. Eleutiherodactylus cram ptop i
4. Eleuliherodactyl us portoricensis 8. Eleutherodactylus antillensis
Introduced.







SCIENTIFIC SURVEY OF PORTO RICO


Eleutherodaclylus britloni
Eleutherodaciylu.s qwighimnanae
Eleu I erodarlylus. rich mondi
Ele tieroduclylu.s imiirolor
Ilem idactyluis nmbouiia
Sphaerodaclylus mua'rolepi.s
A nolis cuvieri
A nolis crislatellus
Anolis gundlachii
Anolis everman ni
Anolis s/rinfulus
Anolis krugi
A nolis pulchellits
Anolis poncensis


23. tC'yclura porloricensis
24. Celestus pleii
25. Ameiva e.rsul
26. Ameivu weetmnorei
27. Amiipiisbmaeian caeca
28. Am phisbaena bakeri
29. Mabuya sloanll
30. Typlilops platycephalus
31. Typhlops roslellatus
32. Epicrates inornaltus
33. Dromicus stahli
34. Alsophis portoricensis
35. Alsophis anlillensis
36. Pseudemys slejiegeri


TI. MONA ISLAND


The fauna of Mona Island, which adds six species to the above list, is
as follows:
1. Eleutherodlctylus nioiiensis G. Mabiuya sloanii
2. Sphaerodactylus macrolepis 7. Typhlops moneiisis
3. Anolis crislalell s 8. Epicrates monensis
4. Cyclura slejneyeri 9. Alsophis variegalus
5. A)meira alboguidlta

III. DESECHEO ISLAND

Desecheo Island is rarely visited. Herpetological specimens were se-
cured by Bowdish in 1901, by Lutz in 1914 and by Danforth in 1926.
The species known are:


1. Anolis crislatellus
2. Ameivr excsul


3. Alsoph is porloricensis


IV. VIEQUES ISLAND

Ten species, all of them identical with Porto Rican forms, are known
from the island of Vieques. These are:


Leptodaclylus albilabris
Eletth erodactylus antiliensis
Sphaerodactylus macrolepis
Anolis cristatellus
Anolis stratidus


6. Anolis pulchellus
7. Anolis curieri
8. Ameiva exrsul
9. Mabnya sloanii
10. Alsophis antillenisis








SCHMIDT. AMPHIBIANS OF PORTO RICO


V. CULEBRA ISLAND

The Culebra fauna lacks Spliaerodlartylis, which has doubtless merely
been overlooked. It adds a Virgin Island form. Dronictns e.rigtns. to
the fauna under consideration. Its species are:
Leplodactylus albilalrris Am eiva e.rsul
Eleuitherodactyl is aitillensis 31abuiaiy sloanii
Anolis cristatellus I)romticus exigitus
Anolis stratuluis Ilsoplit .s autillen2is
Anolis pulch ell/u
VI. CAJA IDE M'EIiTOS ISLAND

Anthony and Goodwin secured four lizards and a snake from this
island during their field-work in 1926. These represent three speie-:
Anolis cristlatellts .lsopJiiS portoricensis
Ameiva wettoirei

HABITAT ASSOCIATIONS ANu FAUNAL SUBDIVISIONS

Porto Rico includes a wide range of habitat conditions, from the ex-
tremely wet mountain rain forest of the Luquillo, where mountain
palms and hardwoods are hung with lianas and draped with moss that
never dries out, to the opposite extreme of aridity on the southwest
corner of the island (near Guanica and Ensenada), where a cactus flora
predominates. Some of the types of habitat, with distinct associations of
reptiles and amphibians, appear to be the following:

I. Northern Coastal Plain (Collections secured from Santurce, Rio
Piedras, Bavamon and Mayagiiez).
II. Coffee Belt, 900-2000 ft. (Collections from Aibonito and Maricao).
IIT. Deforested HIilltops, above 2000 ft. (Collections secured at Aibo-
nito and Maricao).
IV. High Rain Forest, 1200-348.) ft. (Collections secured from El
Yunque, Lnquillo Forest Reserve).
V. Pepino Limestones (Collection from Catailo).
VI. Arid Limestones,. southwestern Porto Rico (Collections from Coamo
Springs, Ensenada and Salinas).

This list is quite inadequate from an ecological standpoint and in it
only II. III and IYV approach the definition of Biotopes, with recogniz-
able Biocoenoses.
Turning first to the distribution of the fauna in Porto Pico itself, a







SCIENTIFIC SURVEY OF PORTO RICO


number of corrections are necessary in the account of the vertical
distribution given by Stejneger. These will be presented in detail be-
low, in the systematic discussion of the species. In general, recent
observations show that altitude in itself has played a relatively small
part in determining the distribution of the fauna. Thus Anolis pul-
chellus, which Stejneger believed to be confined to the coastal plain, be-
low 500 feet, is present at all altitudes, at least up to 2000 feet, in open
fields; and Anolis kruii, for the most part confined to the coffee belt, is
found as far down as Coamo Springs (500 ft. alt.) where the conditions
of moisture and shade are suitable. The species which are abundant at
the lower altitudes (i. e., on the coastal plain) and extend in varying de-
grees into the higher are the following:


Bufo lemtvr*
Leplodactylus albilabris
Eleutherodactylus porioricensis
Eleutherodactylus antillensis
IHemidactylus maboUtia*
Sphaerodactylus macrolepis
Anolis cuvieri
Anolis cristatellus
Anolis stratulus
Anolis pulchellus
An-olis poncensis*


12. Ameira exsul*
13. Ameiva wetmorei*
14. Amphlisboena caeca
15. Mabuya sloanii*
16. Typh lops pla yceph alus*
17. Typhlops rostellatus
18. Epicrates inornatus
19. Dromicuts slahli
20. Alsophis portoricenis.
21. Alsoph is antillensis*
22. P'seudemys stejnegeri*


Of these only nine (marked with an asterisk) are, so far as known,
confined to the coastal plain, or to altitudes below 500 feet. Bufo
marinus may now be added to this list.
The species, on the other hand, which do not occur on the coastal plain
or at least only as stragglers, are the following:
1. Eleutlherodactyltus gryllus 7. Eleutherodactylus unicolor
2. Eleuiherodactylus locusts S. A nolis h,.;n..i ,,
3. Eleuthlerodactylus crampfoni 9. Anolis evermanni
5. Eleuiherodactylus brittoni 10. .1noli. krugi
5. Eleutherodactylus richmondi 11. Celestus pleii
6. Eleutherodaclylus wightmana.e 12. Amphisba.ena baker
Of these Eleutherodactylus crampioni, E. unicolor and E. richmondli
and E. locusius are confined, so far as known, to the peak of El Yunque:
the others are probably most abundant in the coffee belt. Since nearly
two-thirds of the coastal-plain species overlap the coffee belt in dis-








SCHMIDT, AMPHIBIANS OF PORTO RICO


tribution, it seems obvious that the distribution in altitude offers little
basis for a faunal division. The changes due to cultivation, it may be
assumed, have played an important part in the present distribution.
The clearing of lowland forests, for example, has undoubtedly driven
species to the coffee belt and to the residual forests, while the clearing of
the hills has probably afforded access to the higher altitudes in the case
of species originally confined to the more open spaces in the coastal plain.
The coastal-plain fauna, however, is not a homogeneous one. Anolis
poncensis and Ameiva we/morei and possibly Alsophis antillensis are
confined to the arid or semiarid southwestern part of the island, and
Eleutherodactylus untilleasis, Anolis cuvieri, Typhlops platyceplhalus and
Typhlops rostella/us have not been recorded from that part of the island.
Anolis poncensis and Aimeira wetmorei are two of the most peculiar and
striking species in the entire fauna, the latter being more closely related
to species in Hispaniola and St. Croix than to other Porto Rican forms.
I propose, then, to divide Porto Rico faunally into a humid district,
comprising the greater part of the island, characterized by the presence
of Eleutlhorodactylu.s antillensis, Anolis cuvieri and Typhlops rostellalus
(besides the species of Eleueherodaclylus confined to El Yunque) ; and
an arid district, including the southwestern corner, characterized by the
presence of Anolis poncensis and Ameira wetmorei. Various cacti form
the most characteristic element in the flora of the arid district (Plate I),
while the humid district was probably originally a forested area (Plates
II and III), bordered by open spaces along the coast.
The contrast in habitat conditions between the arid area to the south-
west and the dripping cloud forest of the Luquillo is extreme. The
cloud forest affords ideal conditions for the tree frogs, and these are
extraordinarily abundant in the moister belt above 1200 feet altitude.
Thlie amphibian chorus in the rain forest on El Yunque is the most
extraordinary I have heard. As one stands at the Forester's Cabin,
at about 1300 ft. altitude, a roar of sound comes from thie wooded
ravine adjoining, and from the slopes above, making a veritable Babel of
frog notes. One by one the individual voices can be dissociated from the
general confusion. Those of Leptodaclylus albilabris and Eleutherodac-
tylas portoricensis, become separated first, since these are already familiar
from the first night in Porto Rico. E. portorice.nsis here appears to have
added several variations to its lowland notes, but in general its voice
proves readily distinguishable. Next to these, the most insistent ele-
ment in the chorus is a rapid click-clicking not unlike that of a tele-
graphic instrument, with a very insect-like quality. This proves to be







NCIENTIFIC SUNVI-IEY OF PI'ORTO RICO


the note of the tiny Eleutherodorlylus yryll.is, and it was undoubtedly
this note which Stejneger ascribed to the young of E. porloricesis. This
clicking note comes also from the lower branches of the trees, probably
up to a height of twenty feet. A fourth note, carefully run down, proves
to be that of a large, green, long-horned grasshopper, and to the sur-
prise of the collector another succession of sounds, even more character-
istically grasshopper-like, beginning with a shrill prolonged note and
ending with a series of clicks, proves to issue from the distended throat
of still another Eleuthlerodaclytluo. Directing the attention, now, as
much as possible away from the known elements of the chorus, one may
distinguish a strikingly different element. A sad little series of whis-
tles descending in the scale and becoming successively fainter proves to
belong to a very distinct species of small Eleutherodactylu." (E. wuight-
nmanag), which sits on the ground or on the lower leaves of plants, and is
certainly a most difficult species to discern, even when it is singing a foot
away from the collector's ear. Another tiny species has a slow clicking
note,-the sixth to be distinguished. There is still an undifferentiated
chorus awaiting investigation, and three species of tree frogs (E. rich-
moidi, E. unicolor and E. crauiponoi) are known from El Yunque,
whose notes I did not trace.
In the arid southwestern section there is no such wealth of amlphib-
ians and, while this is obviously dcue to the lack of moisture and hence
is primarily an ecological difference, the differentiation of very distinct
species confined to this area bears witness to so long a history of similar
relations between topography and moisture that here habitat conditions
have dominated the faunal history. The fact that this section of Porto
Rico appears to be intimately related to the island of St. Croix, figures
in my argument below on the relations of the faunae.

ORIGIN AND RELATIONS OF THE P1ORTO RICAN HEII'PETOIOGICAIL FATTiA
I. THE WEST INDIAN FAUNA
The origin of the West Indian fauna, specifically of the Greater An-
tillean fauna, has been a controversial topic among zoogeographers for a
generation. Arguing from herpetological evidence, Stejneger (1904)
and Barbour (1910, 1914, 1916) have maintained that the fauna is de-
rived from the mainland by migration over land connections, and An-
thony (1918) supports the same view from the standpoint of mammal-
ogy. Matthew (1915, 1919) has been the chief exponent of the alter-
native theory that the Antilles have received their fauna through fortui-
tous dispersal without such connection.








RSHMIDT, AMPHII'IAI.NS OF PORTO RICO


Anthony (1925), in summarizing the evidence from the mammalian
fauna in an earlier volume of the present series, adopts a modified
form of the "land-connection" hypothesis, and Matthew himself (1919).
has agreed that the Greater Antillean islands may at some time have
been united. The West Indian amphibians and reptiles appear to me to
afford evidence supporting Anthony's conclusions, at least in a general
way.
Comparison between the distribution of amphibians and reptiles and
the distribution of mammals is made difficult by the much greater age
of amphibian and reptile stocks. The arrival of the bulk of the West
Indian reptile fauna may be contemporary with that of the earliest of
the mammals, the insectivores, whose mammalian contemporaries are ex-
tinet. Reptilian distribution frequently affords clues to pre-mammalian
faunal history. Thus Madagascar and New Zealand may be allowed to
be oceanic islands so far as their mammalian faunae are concerned,
while their Pre-Tertiary contacts with continental faunae are reflected in
their amphibians and reptiles.
From a general review of the distribution of the reptiles L am con-
vinced that they support the general theses of Matthew regarding the
trend of dispersal from IIolarctic centers and the want of evidence for
Antarctic connections. I am equally convinced that reptilian dis-
tribution fails to support some of his secondary theses, especially with
regard to the oceanic nature of the faunae of Madagascar and the West
Indies. It is embarrassing to be so thoroughly an eclectic zoogeographer,
and one finds oneself exposed to the fire of both schools.
My own general conclusions with regard to the West Indian fauna,
based primarily on the herpetological evidence, are:
1. That the Greater Antilles received their fauna from Central Amnier-
ica at a time so early that the continental fauna has subsequently under-
gone great changes, probably in Eocene or even in Pre-Tertiary time.
2. That the Greater Antillean fauna gives us a somewhat obscure
representation of this earlier Central American fauna, most of which,
in accordance with Matthew's general hypothesis, has moved on to South
America.
3. That there has been a union of the larger islands during part of
their existence, which has produced the uniformities in their faunae.
4. That the Lesser Antillean fauna is derived from South America,
that it is a genuinely fortuitous one and that no land-bridge has existed
through this chain in Tertiary time.
By way of general review of the Greater Antillean herpetological
fauna, I have drawn up a list of the genera in tabular form.









14 SNCENTIFIC SURVEY OF PORTO RICO


Number of species native on the Greater
Antillean Islands
List of Genera

ub Jmi His- Porto Virgin
Cuba Jamaica paniola Rico Islands


AM PHIBIANS
1. B ufo *t..................... 5 0 1 1 1
2. H yla *t..................... 1 4 4 0 0
3. Leptodactylus* f .............. 0 0 1 1 1
4. Eleutherodactylus *f ......... 16 8 9 10 2
5. Sminlhillus t ............. 1 0 0 0 0
REPTILES
I. Gonatodes *t ................. 1 1 1 0 0
2. Sphaerodactylus *f............ 5 6 5 1 1
3. Hemidactylus f ............. I 1 2 1 1
4. Aristelliger ............. ... 1 1 1 0 0
5. Tarentola ................... 1 0 0 0 0
6. Thecadactylus *f ............. 0 0 0 0 1
7. Anolis .................... 25 6 13 8 6
8. Norops *t. ................... 1 0 0 0 0
9. Deiroptyx ................... 1 0 0 0 0
10. Chamaeleolis ........ ....... 1 0 0 0 0
11. Chamaelinorops .............. 0 0 2 0 0
12. Xiphocercus ................. 0 1 0 0 0
13. Iguana ..................... 0 0 0 0 1
14. Cyclura *t ................... 1 1 3 1 1
15. Leiocephal s t .............. 5 0 8 0 0
16. Hispaniolus................ .. 0 0 1 0 0
17. Celestus*................... 1 2 3 1 0
18. Sauresia .................... 0 0 1 0 0
19. W etmorena.................. 0 0 1 0 0
20. Cricosaura .................. 1 0 0 0 0
21. Ameiva*t................... 1 1 8 3 2
22. Amphisbaenat............... 1 0 3 2 1
23. Cadea ..................... 2 0 0 0 0
24. M abuya*ft.................. 0 1 1 1 1
25. Typhlopst ................. 1 1 2 3 1
26. Tropidophis*t ............... 4 1 2 0 0
27. Epicratest .................. 1 1 3 2 0
28. Tretanorhinus* .............. 2 0 0 0 0
29. Arrhyton.................... 3 0 0 0 0
30. Alsophist ................... 3 0 3 3 2
31. Dromicus ?f................. 2 2 5 2 1
32. Uromacer ................... 0 0 5 0 0
33. Hypsirynchus ............... 0 0 1 0 0
34. Ialtris ...................... 0 0 1 0 0
35. Pseudemys* ................. 1 1 1 1 0
36. Crocodylus*f ................ 2 1 1 0 0

Total species.............. 91 40 92 41 23

Total genera.. ......... 29 18 29 16 15
Endemic genera........... 5 1 7 0 0

Non endemic genera not
found in other islands.... 5 0 0 0 2


* Central American.


I South American,








SCHMIDT, AMPHIBIANS OF PORTO RICO


The number of species in this table is somewhat unsatisfactory for
comparison on account of the inclusion of vicarious forms from out-
lying islands-the Cayman Islands with Cuba; Tortuga, Gonaives,
Navassa and Beata with Hispaniola, and Mona with Porto Rico.
The Amphibians and Reptiles of the Greater Antilles represent 41
genera. Two of these, Iguana and Thecadactylus enter the region only
in the Virgin Islands, and are present in the Lesser Antilles. They are
consequently an alien element in the fauna, the more so as they are not
specifically differentiated; it is extremely likely that Iguana was intro-
duced by the Indians in the course of their wanderings, while the gecko
is probably fortuitous through non-human agencies. A third genus,
Tarentola, is represented only in Cuba and is otherwise African, spe-
cifically Mediterranean, in distribution. This still more alien form is
well differentiated from its congeners and represents one of the most
curious of genuinely discontinuous distributions. I suppose it to be an
ancient "flotsam-jetsam" arrival.
Of the remaining 37 genera, 14 are endemic: 11 are generally dis-
tributed on the four larger islands, and 20 are represented on three or
more of the islands. It is a curious fact that the endemic genera, with
the exception of Cyclura, are confined to single islands, and thus do not
contribute to the hypothesis of a former union. The 20 more widely
distributed genera, however, all have vicariating forms from island to
island, and a number of sections of genera, such as the giant Anoles.
come near to being widely distributed endemic genera, like Cyclura.
The endemic forms are chiefly minor end-stages or divergent branches
which have arisen by local evolution, such as Chamaeleolis, Deiroptyx.
Chamaelinorops, Xiphocercus, Hispaniolus, Sa:aresia, Welmorena and
Arrhylon. A few, however, are plainly relict forms, notably the Xan-
tusid Cricosaura, the Iguanid Cyclura, the IIispaniolan snakes Uromacer
and Ialtris, the Brachycephalid frog Sminthillis. Five genera, Leplo-
daclylus, Sminthillus, Norops, Leiocephalus and Trelanorhinus, are
neither endemic nor widely distributed, and this is a very heterogenous
list, with no appreciable parallelism in distribution.
Eighteen genera occur both in the Greater Antilles and Central
America, but 14 of these are likewise represented in South America,
and these, with the 7 genera common to South America and the
larger West Indian islands but absent in Central America, make the
faunal relation with South America appreciably more intimate than
with Central America. This very fact seems to me to accord best with
the theory of the Central American origin of the fauna, on the supposi-








SCIENTIFIC SURVEY OF PORTO RICO


tion that the South American fauna is mainly of northern origin, as
pointed out by Matthew in his general scheme of dispersal.
The degree of differentiation between the continental and West Indian
representatives varies greatly, and at first glance appears to indicate
varying ages of origin. Some of this variation, however, may be due
to other factors than time of separation. Such an archaic-looking relict
as Crirosaura, widely distinct from its continental allies, may perhaps
represent about the same amount of evolution as has occurred in Anolis
and its derived genera, the difference being the contrast between a de-
clining group and an expanding one. The crocodiles, on the other hand,
seem to belong to quite different invasions, C. rhownbifer and moreletii
being assignable to an earlier arrival, their ranges now entirely circum-
scribed by that of the modern wide-ranging, semi-marine Crocodylus
ac1uus, whose wide distribution evidently has little bearing on the prob-
lem of land connections.
In a more detailed discussion of the genera I shall try to show that
the faunal picture presented accords with a derivation from Central
America at an early date, on the hypothesis of a southward trend in the
migrations of the world as a whole, and that it is direct faunal relations
with Central America, such as that of the Xantusiidae, which require
explanation rather than the discontinuity in range of A mphisbaena or
Leiocephalus.
Of the genera of Amphibians, Bufo, Hyla, Eleutherodactylus and
Leptodwutylus have a wide Neotropical distribution. The anomalous
nature of the distribution of Leptodactylus will be discussed below.
Smintlhillus has a single Cuban species, and two others, Peruvian and
Brazilian. have since been described. The discovery of additional species
in this genus (originally described as monotypic) contributes to the
likelihood that it is a natural group.
Among the reptiles, geckos are notable for erratic distributions, though
when critically examined their ranges are often found to be closely
parallel to those of other groups. The Antillean geckos, however, are
really heterogeneous in distribution. T.i,- ,.*i,., and Thecadactyluys have
already been mentioned. Gonaaodes is widely distributed in Central and
South America, apparently ranging into the Antilles from the west.
Sphlaerodactylus has a wide neotropical distribution, but its wealth of
Antillean species distinguishes it as an autochthonous genus, and its
development is very like the other characteristically West Indian forms.
such as Aneiva, Dromicus or Eleatherodaclyluo. Hentidactilus, with IIH.
manbonia on all the islands, appears to be a house-gecko. and humna'n








C('II'ItT. AMJIP'IIIAN OF PORTO RICO


agency may well have played a part in its distribution. It is somewhat
remarkable that the genus llemidac/gls" is unknown in Central Amer-
ica. I am not at all convinced that the African geckos commonly re-
ferred to maboula are con-specific with the Antillean form. The East
African and Madagascan species does not seem to me to be identical
even with the West African one! HIemidactylus brookii, on the other
hand, in Hispaniola, would appear to be an African form introduced
by the slave-trade. A ritrelli'yer is confined to Central America, Jamaica
and Hispaniola. It is included in my list as Cuban because it reaches
the Cayman Islands, whose faunal affinity is primarily Cuban. On the
coast of Yucatan this species is characteristic of the fringe of cays, and its
occurrence in the West Indies offers no anomaly.
The Iguanid genera include the monotypic and endemic Deiroply.r,
Chiamaeleolis, Chamaelintorops and Xiphocercus, Iguatna, already men-
tioned, and Anoli., Norops, Cyclura, Leiocephalus and li.spati olus.
Norops seems to be a more primitive form than Anolls, with three con-
tinental species, and is a declining group in contrast with the expanding
A nolis. The Cuban species is thus plainly a relict. Leiocephalus, with
a number of species in Cuba and Hispaniola, is otherwise best developed
in western South America, and is absent in Central America. I regard
this also as a reliet distribution, but of a group that is holding its own.
Cyclira is even more interesting. The curious "eombs" on its toes,
though rather a trivial character, quite definitely ally its species more
closely to the Galapagan Conolophuis and Atotdlyrhy/ichus (and the other
Pacific genus as well, the Fijian Brachylophus) than to the Central
American Cltenoitura. (tenosaura extends southward as a wedge sepa-
rating these allied forms, and I hase endeavored elsewhere* to show
that the Ctetosaura have spread southward from the great Southwestern
shield in North America. Lelocephutlus lends itself to this interpreta-
tion if it be visualized as retreating before more advanced Iguanid
genera, such as Sceloporus. Anolis, in the full flower of expansion, ob-
scures distributional argument by its wealth of forms and closely-knit
ranges. The only species of Anolis that is supposed to be common to
Central America and Cuba is A sagrei, an inhabitant, like Arildelligcr,
of the off-shore cays in the Bay of Ionudura,. The endemic genera re-
quire no comment except that Aipth ocercu.s is represented in Colombia
by a related or parallel form.
The Anguidae are represented in all four islands by Celestus. Two
additional genera. `"auresia and WlVemorena, monotypic "end-stage"
1922. Bull. Amer. 3Mis. Nat. Hlst., XLVI, p. ip. 1.








SCIENTIFIC SURVEY OF PORTO RICO


forms, are confined to Hispaniola. Celestus also occurs in Central
America and its close ally, Diploglossus, is found in both Central and
South America. Celestus and Diploglossus are plainly primitive genera,
and the modern representatives of the family, the plated lizards (Gerr-
honotus), have the same spatial relations with them as exist between
Ctenosaura and Cyclura, or Sceloporus and Leiocephalus.
The Xantusiidae are a declining group composed of the North Ameri-
can genus Xantusia, the Central American Lepidophyma and the Cuban
Cricosaura. This distribution is not in accord with the above-cited
southward migrations, but this is a recurrent anomaly which requires
a modification of the Matthewsian hypothesis of the dispersal of primi-
tive forms. It must be recognized that evolution in the direction of
habitat restriction may strictly parallel an evolution in which the primi-
tive forms become peripheral by retreat in space. This is an obvious
phenomenon among the Xanlusids, which inhabit areas adjacent to what
I have regarded as the probable center of dispersal of American lizards,
but are plainly relicts among more modern and progressive forms. The
species of Xantusia are curiously restricted as to habitat-X. vigilis by
its association with the Yucca, X. henshliaw by its rock-dwelling habit-
while both are doubtless nocturnal, as is Lepidophyma. The mainland
Xantusids have retreated owing to habitat restriction, while the Cuban
genus represents the other alternative, that of actual retreat, and appears
as a true relict, though also rigidly confined to a single habitat.
The case is directly comparable to that of the Central African lemurs,
which escape their modern competitors by their nocturnal habits, while
the Madagascan lemurs have survived through actual migration and
the timely separation of their retreat.
The Teiidae are represented only by A meira, though the West Indian
species are divisible into two rather distinct sections. Ameivas are
widely distributed on the South and Central American mainland, but the
continental species are fewer than the West Indian. I suspect that the
genus COnemidophoruis hears the same relation to Ameira as Sceloporus
does to Leiocephalus, namely, that it is a more modern group of species,
with Ameiva more or less in retreat.
The Amphisbaenidae are well represented in the Antilles, with both
Cadea and A mphisbaena in Cuba and Amp iisbaena extending out to the
Virgin Islands. Except for Bipes, which is present in west Mexico,
the family is wanting in Central America, and the Antillean forms are
thus relicts of a former type of dispersal. The evidence for the
southward migration of the Amphisbaenians seems to me ample, even








SCHMIDT, AMPHIBIANS OF PORTO RICO


without the direct evidence of the Oligocene fossil forms. Rhincura,
confined to Florida, is quite obviously one of the many curious forms
accumulated in the southeastern United States as a result of divergent
migration from holarctic dispersal centers. The nearest relative of
Cade seems to be Venezuelan, while A mphisbaeiea itself is well repre-
sented almost throughout South America.
The only Scincoid genus is Mabtya. generally distributed in the
tropics of the world, but nowhere speciating in the Americas as its does
in the Old World. Its range in both hemispheres is nearly exclusive of
that of the more northern and obviously more recent Eumeces.
Among snakes the Typhlopidae afford no especial evidence of faunal
relation. The Antillean Typhlops luminbricalis was long supposed to be
a widespread species occurring also in South America. Cochran (19124)
and I (1920) have brought the distribution of the West Indian forms
into harmony with that of other groups. The most notably primitive
genus, Anomalepis, is Central and South American (or at least Pana-
manian and Peruvian), and not Antillean.
The Boidae are represented by two genera. Elpicrates has a species on
each of the larger islands and has split into three species in Hispaniola,
with a separate species on Mona Island and another distinct form in the
Bahamas (confined to Turk's Islands). Tropidophis fails to reach
Porto Rico, and its principal radiation occurs in Cuba. Epicrates is
wanting in northern Central America, but it reappears in South Amer-
ica. Tropidophis is said to have both South and Central American allies,
but they are little known.
The relationship of the Colubrine genera are vague, but their nearest
allies seem to be South American, with the exception of Tretior, i aus,
which is found in Cuba and Central America.
Pseudemys, the single genus of fresh-water turtles, is quite as easily
derivable from the Central American representatives as from the Florid-
ian, and the existence of insular differentiation, which I am able to show
for the Porto Rican specimens, makes it unnecessary to regard Pseudemys
as a strictly recent arrival. The absence of other fresh-water turtles is
highly remarkable, in view of the ancient character and great diversity
of the American turtle fauna. It is no less anomalous to find in Cuba
a fossil Testudo related to the Galapagan species, though its presence
adds to the faunal relations between the Antilles and the Galapagos.
Crocodylus, finally, adds a distinctively Central American form to the
West Indian fauna. The broad-snouted Cuban Crocodylu.s rhombifer
is directly allied to C. moreletii of the adjacent parts of Mexico and the
Yucatan peninsula. The wide-ranging, undifferentiated C. acutus








SCIENTIFIC SURVEY OF PORTO RICO


floods over the ranges of these earlier forms. Crocodiles do not range
beyond the Orinoco hasin in South America, and evidently are more
recent arrivals than the caimans or the alligators.
In support of my proposition (2) above, I have contrasted the distri-
bution of such a group as Sceloporus, an essentially modern genus. with
that of a more ancient Iguanid genus, Leiocephalus. Sceloporus is
essentially Sonoran, with a wealth of North American species, and a
broad overflow into Central America. Lelocephalus is West Indian
and South American. Allowing for discrepancies and irregularities such
as those I have discussed for the Xantusiidw. the list of s-uch pairs of
genera is impressive:
Ancient. West Indian Modern, Sonoran
Leiocephalots Sceloporus
('yclura Ctenosaura
('elestus Gerrlihoolus
.1I eira Cnemtidoph orus
Mubwu/a Eum ece.

Cnemidophorus, among the genera listed as Sonoran, ranges widely
into Sonth America. Otherwise its development is so closely similar to
that, of the other Sonoran genera that I am disposed to search for an
explanation of this anomaly rather than remove it from the Modern,
onloran list.
I am fully convinced that the fauna of the Greater Antilles reached
these islands from Central America. and that the majority of the endemic
forms represent a nearly contemporary faunal invasion. That an actual
landbridge existed over which the migration took place, is my somewhat
more hesitant belief. The existence of mammals and amphibians, even as
a depauperate fauna, is evidence in favor of continental connection. The .
amphibian and reptile faumna exhibits a relatively greater diversity
than does the minannalian. The sixteen families represented are:
Amphibians
Blu onidae llylidae B]rachyceephalidae
TReptiles
(Gekkonidlae Amphisbaenidae Boigidae
]gnanidate Scincidae Einvdidae
Anguidae Typhlopidae Crocodilidae
Xantusiidae Boidae
Teiilne Colubridae








SC'HMII117, AMHI'/IIAHN OF I'ORTO R1ICO


The Central American 'anuna has thirty-two families, but several
of these are isolated groups which could scarcely be expected in the West
Indies.-the IHelodermatidai and Xenosauride, for example. Others are
obviously more recent arrivals and for this reason their presence is not to
be expected; instances in point are tile Ranidae. Plethodontidae and Cro-
talidae. The disproportion between the continental and Antillean faune
in number of families is accordingly much reduced, perhaps about 26:16.
It is a striking and important fact that the South American fauna is
actually poorer in families of amphibians and reptiles than the Central
American by four or five. If the Chelydridw. Crocodilida, and I'letho-
dontide, which are essentially Central American andI only enter South
America at the northwest, are also excluded, tile genuinely South
American families number only twenty-five.
If the date of the supposed continental connection of the West Indies
be placed at the close of the Mesozoic. the relative wealth of amphibians
and reptiles and the poverty in tuntmnals are comniletely explained. ('n-
fortunately, a connection so early in geological history doee not account
for the more recent members of the mammalian fauna, for which a Mio-
cene date of arrival is indicated. The two families of insectivores
agree with the reptiles as to-'elarly date of entry, while the remaining
manmmnals appear to represent at least two later immnigration.s. One is
tempted to suppose a very early continental connection for amphibians
and reptiles, insectivores, etc.. and to recognize Matthew's argument that
the remaining mamiials are accidental. Geological conclusions based on
zoogeographic evidence so fragmentary and contradictory are evidently
of little real value.
One set of conclusions, however, from a general consideration of the
fauna, seems well founded. This is my proposition (3), that the larger
islands were connected at an early stage in the development of their
fauna: that they have subsequently been separated, more probably )by
block-faulting than by any great change of level, that Porto Hice and
the Virgin Islands were the last to be cut off, and that the V-irgin
Islands were connected with Porto Rico as recently as the Pleistocene.
The evidence of long isolation of the three western islands is plainly to
be seen in the independent radiation which has taken place in the
elements of their fauna,. Itifo lias evolved 5 species in Cuba. while
Jamaica with no native toad, and 11lispaniola with a single Tlufo, have
each produced 4 species of H/lIh, independently. in the opinion of Dunn,
who has lately examined the Jamaican species in detail. Donn's concl-
sions are somewhat hesitatingly accepted by Noble (1!12). Whether or








SCIENTIFIC SURVEY OF PORTO RICO


not the radiation in lIlla, has been independent, the fact that both
Jamaica and Cuba have only a single species of A meira while Hispaniola
has 8, and 4 on the main island alone, certainly illustrates an inde-
pendent evolution. Long separation is equally evident in the wealth of
Cuban A nolis with no less than 3 related genera represented, Norops,
Chamceleolis and Deiroply.r, equalling the number of species on the other
three islands together. The Jamaican fauna, poor in some genera, has
no less than 6 well-established species of Spha'rodaclylus, surely a sharp
contrast with the 2 in all Central America!
Among the snake genera, Tropidophis has developed 4 species in
Cuba, while Epicrates has 3 in Hispaniola, and an extra species on
Mona. The IIispaniolan Dromicus, Alsophiis and Uromacer fall in line
with the other genera. The total impression of the herpetological fauna
is plainly one of a fundamental unity, obscured only by the long exolu-
tion during subsequent isolation.
The fauna of the Lesser Antilles has been effectively contrasted by
Anthony with that of the larger islands. His argument from the mam-
mals that the aniiial population of the Lesser Antilles is fortuitous
from South America and of relatively recent origin agrees exactly with
my impression based upon the herpetological fauna. The fauna of
Trinidad itself is far from rich in comparison with that of the mainland.
Its reptiles and amphibians amount to about 80 species-almost all
of them specifically identical with those of northern South America.
This South American fauna disappears rather in proportion to distance
from the mainland than in relation to size of landmass, for Tobago has
24 species with little endemism, Grenada 17 with about 4 endemic forms,
St. Vincent 10 with G endemic. In the next four islands the fauna
ranges only from 10 to 14 species, with 5 to 10 endemic forms. En-
demic forms in the whole chain are very slightly differentiated from their
very obvious relatives. The species may be grouped as mainland forms,
with a haphazard distribution on the islands, endemic species of main-
land genera, slightly differentiated, and endemic species of genera which
range throughout the chain with vicariating forms from island to island.
There is little or no "radiation," which is so marked a characteristic of
the Greater Antilles. Examples of the haphazard distribution are
afforded by Leplodactylius and Iguana, probably transported by the
Indians as food animals, and by the snakes in general, though the faunw
in question may be imperfectly known, Leptotyphlops bilineatlus, for
example, occurring on Barbados and St. Lucia, Cloelia clelia on Grenada,
St. Lucia and Dominica, although not recorded from the intervening








SCHMIDT, AMPHIBIANS OF PORTO RIUO


islands, St. Vincent and Martinique. The fer-de-lance skips Tobago,
Grenada and St. Vincent, to appear only on St. Lucia and Martinique.
The northern group of twelve small islands, from Anguilla to Mont-
serrat, has a fauna impoverished in genera, but rich in endemic species.
Leptodautylis peni ulactylus and Iguana iguana appear to represent Pre-
Columbian introduction by natives. Eleutherodactylus martinicen,.is is
recorded from five islands. Its status requires re-investigation. Ty-
phlops is known from St. Kitts and Antigua, the species doubtless unde-
scribed. Leplotyphilops albifrons is reported only front Antigua. Its
wide range and apparently haphazard arrangement seems to indicate a
facility for fortuitous dispersal. Thecadactyllus rapicaudus, widespread in
the Lesser Antilles, is recorded from five of the northern islands. These
irregular distributions contrast strongly with 3 endemic species of
Sphaeroductylus on three islands, 7 Anolis on nine islands, and 8 A mei-
va on ten islands. This portion of the fauna, which perhaps should in-
clude Alsophis, appears to represent an older nucleus, and I interpret its
relations as representative of the uniformity of a typically oceanic fauna
plus endemism induced by a considerable lapse of time.
The most obvious differences theoretically to be expected between
continental and oceanic insular fauna are (1) heterogeneity and (2)
impoverishment in the oceanic islands. The presence of relict forms
seems to me to be strong evidence of a land connection remote in time.
Impoverishment may obviously occur in a continental fauna by extinc-
tion; and "fortuitous dispersal" may act as a screen allowing only cer-
tain forms to pass, so that extreme uniformity of fauna, instead of
heterogeneity, may be a result of truly oceanic dispersal, as is to be seen
in Polynesia, where island after island is inhabited by the same five
species of lizards. Such uniformity is complicated by the age of the
islands. It is thus curiously difficult to frame criteria whereby an in-
sular fauna derived from land connection may be distinguished from one
produced by "fortuitous dispersal."

II. THE PORTO RICAN AND VIRGIN ISLAND FAUNA

Turning to the more detailed consideration of the Porto Rican fauna,
it is interesting to note that important advances have been made in our
knowledge of the amphibians and reptiles of this area since Stejneger's
discussion of their origin and relations in 1904. The herpetological dis-
coveries bearing directly on this problem have been (1) the description
of Ameiva wetmorei, (2) the finding of Bufo and Cyclura on the outer
Virgin Islands, (3) the discovery of fossil remains of Cyclura on St.







NCIN'1FICSun PLj'Y OF PORiTO MICO


Thomas and Porto Rico, (4) additions to the fauna of Hispaniola,
especially the discovery of LeptodaclyIus dominicensis, (5) additions to
the fauna of Porto Rico, especially Alsophis antillensis and Eleutliero-
dactylus anlillensi.s. which ally it more intimately to the fanna of the
Virgin Islands, and (6) the elucidation of the relations of the Greater
Antillean Typhilops. All of this new information has tended to empha-
size the essentially Greater Antillean character of the Porto Rican fauna.
Stejneger divides the Porto Rican herpetological fauna into South
American and Central American elements, including in the former the
genera -A mrira. A mph.isbaena, Typhllops, Al sophis and Dromicius. These
genera are all represented in lisipaniola, and their immediate presence
in Porto Rico is amply accounted for by a union with Hispaniola. I
have endeavored to show shaove that this apparent relation of the West
Indian fauna with the South American is best explained by a land
connection with Central America, when the time relations and larger
outlines of faunal migration are considered.
Vaughan (1919, Bull. TU. S. Nation. Mus., No. 103, pp. 547-612) has
given an excellent r6sumi( of the geological history of the West Indian
area as far as known. In advocating the existence of former land con-
nections with South and Central America, his paper cuts incisively into
the more speenulative maze of zoogeographic controversy. The only flaw
is the fact that lie appears to base his conclusions ii part on zoogeo-
graphical data (p. !lO), whereas 1 should like to accept them as a basis
for zoogeography. For the present purpose however.-an examination
of the immediate origin of the Porto Ricean reptile and amphibian
fauna,-the outline of the geologic history advanced by Vaughan is
highly satisfactory, and it is possible to crystalize conclusions on the
relations between Porto Rico and thle Virgin Islands with each other
and with Hispaniola into a more definite statement than has hitherto
been possible.
According to Vaughan's physiographic history of the area in question.
the Greater Antilles were joined to one another in late Miocene time, the
resulting landmass including Porto Pico and the Virgin Islands as its
easternmost extension. The scanty zoogeographic ties between the Vir-
gin Islands and the Lesser Antilles exclude the presence of a contemipo-
raneous land bridge to South America, or at least the continuation of any
such bridge for a time commensurable with that of the union of the
Greater Antilles. During the period of this uplift, the genera of rep-
tiles and amphibians which may properly be regarded as "Greater
Antillean" (through presence on three or more of the larger islands)
acquired their distribution.








SC'HMIDT, A.1/'IIlIANLVS O/' l'OItTO Ri'")


]n order to connect the Virgin Islands with Porto Rico, no great
degree of emergence is necessary, as they are separated by water not ex-
ceeding twenty fathoms in depth. St. Croix is included in the same way,
with no greater amount of emergence, because its present separation by
greater depths of water is believed to be due to fanlting. The water
between Porto Rico and Santo Domingo is much deeper (reaching 318
fathoms), but still shallow in comparison with the depths to the north
and south. Even for this connection, however, the amount of emergence
necessary is no greater, for there is important evidence of faulting,
as I have shown elsewhere (1926). 'he earthquakes of October, 1918.
which caused great damage to the cities of Mayagiiez and Agnadilla, on
the west coast of Porto Rico., were probably caused by adjustments in
this faulted area. The sharp trmncation of the eastern end of Santo
Domingo doubtless represents another fault line. Point Jrignero and
Desecheo Island appear to represent the older period of mountain-mak-
ing of the general Hlispaniolan-Porto Rican axis, (i. e., an Eocene or
Lower Oligocene connection) while Mona Island, almost exactly half
way between the southwest corner of Porto Rico and Saona Island. and
topographically almost exactly similar to Saona, may be a remnant of
the Upper Miocene (or later ?) land bridge itself. The rapid under-
cutting of the north and east sides of Mona now in progress indicates a
considerable recent reduction of its area. The date of the faulting which
separated Porto Rico from Santo Domingo is placed in the Pliocene by
Vaughan (p. (11)). and the separation of St. Croix from Porto Rico
probably took place during the same period, but perhaps at a later date.
It is likely that Porto Rico and the remaining Virgin Islands were
separated by a submergence in the Pliocene, but they were reunited in
the Pleistocene, perhaps by the withdrawal of the water for the conti-
nental ice sheets, to form a greater r Porto Rico" to which the common
fauna of the islands now separated corresponds. The present configura-
tion of Porto Rico and the Virgin Islands is (geologically) extremely
recent. The very evident peneplanation of the greater part of the
mountain area at a height between 1500 and 2000 feet appears to date
at least from the early Tertiary, and implies long-continued existence as
a land area.
The interpretation of the existing faunal relations of Porto Rico, in
the light of the geological hypothesis, becomes relatively simple. The
fauna of the Virgin Islands stands in the same relation to the Porto
Rican as does that of Porto Rico to the IIispaniolan. The degree of dif-
ference in each case corresponds to the relative length of time since their








SCIENTIFIC SURVEY OF PORTO RICO


respective separations, and the degree of impoverishment to their rela-
tive areas.
The fauna of the Virgin Islands consists of 22 species, of which 2,
Iguana iguana rhinolopha and Thecadactyluis rapicaudus, belong to genera
foreign to Porto Rico. The iguana was probably introduced by man;
the gecko probably is a fortuitous arrival. Of the remaining 20 species,
11 are identical with Porto Hican forms:


Leptodactylus albilabris
Eleuth erodaclylus antillensis
Sphaerodactylus macrolepis
liemidactylus mabouia
Anolis cuvieri
A nolis crislatellus


7. Anolis stratulus
8. Anolis pulchellus
9. Ameiva exsul
10. Mabuya sloanii
11. Alsophis antillensis


The remaining 9 species are directly related to Porto Rican species
and are for the most part simply vicariating forms:


Virgin Islands
1. Bufo turpis
2. Eleutherodactylus lentius
3. Anolis acutus
4. Cyclura pin gis
5. Ameiva polops
6. Amiphisbaena fenestrata
7. Typhlops richardii
8. Dromicus exignus
9. Alsophis sancti-crucis


Porto Rico
Bufo lemur
Eleutherodactylus richm ondi
Anolis poncensis (?)
tCyclura portoricensis (?)
Ameiva wetmorei
A nphtisbaena caeca
Typhlops platycephalus
Dro micus stab li
Alsophis portoricensis


Examined more in detail the chief questions which require discussion
are: (1) the impoverishment of the Virgin Island fauna, in which many
species of Porto Rican Eleutheroductylus and Anolis are unrepresented,
while Celestus, Epicrates and Pseudemys are entirely wanting; (2) the
apparently haphazard distribution of Bufo, Anolis cuvieri and Cyclura:
(3) the position of St. Croix in relation to the other islands and Porto
Rico, and (4) the origin of the species common to several of the Virgin
Islands but absent in Porto Rico.
The absence of forms may be explained as original or secondary. The
discovery of the remains of recently extinct Cyclura in both Porto Rico
and St. Thomas, coupled with the presence of living Cyclura on Mona
and Anegada, obviously indicates that in this genus a process of extinc-
tion is taking place. The same factor probably accounts for the isolated








SCHMIDT, AMPHIBIANS OF PORTO RICO


occurrence of a Bufo on Virgin Gorda and of An olis cuvieri on Tortola.
The absence of Celestius and of Anolis gundlachi and Anolis krugi, may
indicate, on the other hand, that these forms never reached the Virgin
Islands. There is no reason to believe that, if the whole land area were
elevated 150 or 200 feet and so reunited, the species which now avoid
the coastal plain in Porto Rico would be able to reach the Virgin Islands.
The fact that of the entire Virgin Island fauna only Eleutherodactylus
lentus is related to "coffee belt" species in Porto Rico indicates that this
factor has probably operated as an important one in the past.
The mere fluctuation in size of these islands has an important influ-
ence on the rain fall and humidity (and evaporation), i. e., the most im-
portant climatic factors affecting the fauna. The complete submer-
gence of an islet would not be necessary to exterminate the greater part
of its fauna, and it is a differential extermination of this nature which I
believe to be the chief cause of the impoverishment of the Virgin Island
fauna, and possibly of the West Indian fauna in general.
St. Croix presents something of a problem. The Amphisbaena fenes-
trata from that island should be compared again with specimens from St.
Thomas, and with A. caeca. Anolis acutus, Ameiva polops and Alsophtis
sancti-crucis are decidedly less closely allied to Porto Rican species than
are the species from St. Thomas and even the outermost of the northern
Virgin Islands. On the other hand, Ameiva polops indicates a relation-
ship with the arid district of Porto Rico. If the "Greater Porto Rico" at
any time included St. Croix, that area must have belonged to the ex-
tended arid district, which influenced distribution in the "Greater Porto
Rico" exactly as it does in the present. The separation of St. Croix in
Pliocene time by faulting (as suggested by Vaughan) doubtless excluded
it from union with Porto Rico in the Pleistocene, while a Pleistocene
(Glacial period) connection of the other Virgin Islands with Porto Rico
seems highly probable.
Three species-Eleutherodactylus lentus, Amphisbae na fenestmraa and
Dromicus exiguus-are common to two or more of the Virgin Islands
and are absent from Porto Rico. Their development may be explained
as due to a differentiation of the fauna of the lower-lying eastern end
of the "Greater Porto Rico," or to differentiation during the hypotheti-
cal Pliocene separation.
The Porto Rican herpetological fauna differs from that of Hispaniola
chiefly in the absence of the following genera.*

Oedipus is excluded from the Hispaniolan fauna pending verification of its occur-
rence. Dunn regards the Haitian origin of Peter's Oedipus infuscatus as mythical.








NCiEXTIFIC SU'i'/EY OF PORT'O RICO


1. Hyla S. Saur sias
2. Gomonodes 9. Tropidophis
3. Arislelliger 10. Uromacer
4. Leiocephalus 11. Hypsirhynchli us
5. Hispaniolus 42. Ialri.s
6. Cha maelinorops 13. Crocodylus
7. IWeC/orena

Seven of these, Chamnaelitiorops. IHispunni/lts, \I'einorein, Sauresia,
Uromacer, HIypsirhynchus and Jal/ris are confined to HIispaniola, while
the other six are found in Cuba and Jamaica and for the most part in
Central America. An extensive impoverishment of the fanna of Porto
Rico is obviously its most conspicuous characteristic. Recent extinction
may well be admitted as a considerable factor in this impoverishment,
in view of the discovery of the remains of an extinct Porto Rican G'yclnra
as well as by analogy with the extinction of the mammalian fauna. This
may be due to two factors, the restriction of habitat formations due to
increased cultivation, and the changes in climate due to past emergence
and submergence. On the other hand. the much greater altitudes of the
mountains of Hispaniola, and the great diversity of habitat conditions of
that island, of which perhaps the most remarkable is the stratification
of the vegetation on the mountains, makes it highly probable that a
number of forms have developed in .situI, and had not acquired a suf-
ficient range before the separation of Porto Rico to reach it, even if the
habitat conditions of the intervening area were not unfavorable. If the
late M1iocene uplift was not extensive, and if Mona Island is a remnant
of the actual land connection via Saona and southwest Porto Rico, the
habitat conditions of the land-bridge must have been such as to prevent
the spread of many forms. It is more difficult to explain the differences
in the development of such genera as S'phaerodactylus, Celestus, Ameiva
and Epicrales, which have several species on Hispaniola and only one on
Porto Rico.
It is possible that IIispaniola was broken up into several islands dur-
ing the 3liocene, as is indicated by the Miocene deposits which compose
the plain between the Central Sierra and the Monte Cristi Range, and
by the "through valley" of the saline lakes to the southwest.
Only two species are common to Porto Rico and Hispaniola, one of
which, Ilemidactylus mabouia, is a house-gecko and plainly fortuitous,
while the other, ,labuya sloanii, requires critical study. I have seen
no Hispaniolan specimens. The number of species which are closely
related on the two islands is large:








SCHMIDT, AMI'HIBLIXS OF PORTO RICO


Porto Rico
Bufo lemur
Leplodactylus albilabris
Eleutherodaclylus porforicensis
Eleutherodactylas rich mondi
Sphaerodartylus macrolepi.x
Anoll. ruv eri
Anolis crislatellus
Anolis pulchellus
tCyclura portoricensis
Celstus plepi
A meira ersul
A mieia Irelmnorei
A wtphisbaena caeca
Typhlops platycephail us
Epicrates inornatus
Dromicus stahli
A lsopl is pororricensis
Pseudemn s stefjnegeri


Hispaniola
BRufo gutturosus
Leplodaclylus don inicensis
Eleutherodactylus auriculaloides
El eutherodactylus wieinlandi
S1iphaerodactylus difficilis
Anolis 'ricordii
Anolis ryboles
Anolis semnilineatus
Cyrclra corn ua
Celestu. sp.
Aneira. ritlipunrltala
A meira lineolata
Amph isbaenm veinlandi
Ty ph lops sp.
Epicrales slriatus
D)rom icus parrifrons
A Isoph is m elanich n us
Pseudemys palustris


The Mona Island species, especially 'vjyclura steji geri and Epinrales
monensi.s, add important links to this relation.
The remaining Porto Pie'an species, mostly Eleuitherodaclylus and
Anoli.s, which are clearly more closely related to other forms in the
Greater Antilles than to South American or Lessor Antillean species.
may be regarded as the individual development of the Greater Antillean
fauna on Porto Lico, whose mountains occupied a relatively isolated
position during any land connection- that may have existed, certainly
since the early Tertiary.
The general conclusion is that the herpetologicnal f'ain of the "Greater
Porto Rico" is simply an impoverished Greater Antillean fauna. Its
resemblances to the fauna of llispaniiiola are due to land connection, the
date of which is placed by geologists in the UTpper Miocene. The difter-
ences Ibetween the Porto Rican and Ilispaniolan faunas are due: (1) to
a process of extinction still continuing; (2) to the isolated position of
Porto Rico at the eastern end of the land mass, the habitat conditions of
the supposed land-bridge being unsuited to the spread of many forms;
(3) to the differentiation of specifically Porto Rican forms, (a) throulih-
out the Tertiary, the mountain.- of Porto Rico being a center of differ-
entiation for autochthonous forms, as I suppose those of Hispaniola to
have been, and (b) during I ost- 'liocene time. since the separation of
Porto Rico from Hispaniola.









SCIENTIFIC SURVEY OF PORTO RICO


SYSTEMATIC ACCOUNT OF THE SPECIES

Class AMPHIBIA

Order SALIENTIA

Family BUFONIDAI1

KEY TO THE GENERA OF PORTO RICAN FROGS AND TOADS

A. No teeth; a barge parotoid gland on each side of the neck; skin very rough
and warty ..................................................... Bufo












o oj \\




FIG. 1.-Head of toad (left) contrasted with Leptodactylus (right). (From Stejneger.)


'IG. 2.-Foot of Leptodactylus (left) con-
trasted with foot of Elentherodartyilux
(right). Compare slender and expanded
lips of digits. (From Stejueger.)


Q


AA. Maxillary and
smooth.


vomnerine teeth present; no parotoid gland; skin relatively


'The Bufonidae in the broad sense of Noble's revision of the families of Salientia
(Noble, 1922. p. 22) includes the family Leptodactylidae, which, defined in contrast
with the Bufonidae in the restricted sense of older authors, will be found in Stejneger.
1904, p. 509.









SCHM1DT, AMPHIBIANS OF PORTO RICO


B. Tips of fingers not dilated. tapering ..................... Leptodactylus
BB. Tips of fingers dilated into adhesive disks ............ Eluthcrodactylus

Bufo Laurenti
KEY TO THE PORT RICAN SPECIES OF TRUE TOADS
A. Head with prominent hony crests; parotoid gland small and rounded-
Bufo lemiur
AA. Head without long lony crests; parotoid glands enormous, subtriangular-
Blrfo Jnmril u1s

Bufo lemur (Cope)
Salm Concho

Text Figs. 3-4.

Peltaphrync letmur Cope, 1868. Proc. Aead. Nat. Sci. Phila., p. 311.
Bufo lemur Stejneger, 1004. Rept. U. S. Nat. Mus., 1902, p. 570, Figs. 1-5.-
Barbour, 1914. Mem. Mus. Comp. Zool., Vol. XLIV, p. 242; 1917, Proc.
Biol. Soc. Wash., Vol. XXX, p. 102.-Schmidt, 1920, Ann. N. Y. Acad. Sci.,
Vol. XXVIII. p. 108.
RBfo (Pcltaphryne) gutturosus Peters, 1876, Mon. Ber. Akad. Wiss. Berlin, p.
709.
Bufo gutttarosus Gundlach, 1881, Anales Soc. Espafi. Hist. Nat., Vol. X, p.
:14.-Stahl, 1882, Fauina Puerto-Rico, p. 71, p. 161.-Garman, 1887, Bull.
Essex Inst., Vol. XIX, p. 16 (part).

The native name, Sapo concho, referring to the ridged head of the
adults of this species, is an appropriate one. Unfortunately, due to the
rarity of the form, it is practically unknown to the majority of Porto
Ricans and the native boys apply the name indifferently to large speci-
mens of the frog-like Leptodactylus or even of the tree frog.
Type locality.-Porto Rico.
Distribution.-Previous to 1919, this species was known from the
north side of Porto Rico, the only exact localities given being Arecibo,
Bayamon and Vega Baja. Its occurrence at Coamo Springs, nearly at
the opposite side of the island, proves that it is widely distributed. Its
rarity is perhaps due to the mongoose. The close relations of Bufo
turpis Barbour, from Virgin Gorda (British Virgin Islands), with Bufo
lemur constitute important evidence for the close faunal affinity of the
Virgin group with Porto Rico. The relations with the Hispaniolan Bufo
guottrosus are evidently close, and the three species gutturosus, lenur and
turpis are much more closely allied among themselves than any one of
them is to Bufo peltacephalus of Cuba, although peltacephalus must be
considered their Cuban representative.
Specimens collected.-5: Coamo Springs.








SCIENTIFIC SURVEY OF PORTO RICO


Diagnosis.-A true toad with rough skin, rounded parotoid glands on
the shoulders, and high bony ridges on the head.
Original descriplion.-"This is a toad of singular aspect, owing to
the extraordinary development of the bony crests of the cranium and
the large size of its eyes.
"-The muzzle is short and very much flattened, projecting much beyond
the mouth. The upper lip forms indeed a strongly projecting bony rim
all round the mouth. Loreal region very concave, canthus concave and
very close together. The superciliary crests are extraordinarily elevated,
having an arched outline, and descending steeply to the loreal region.
It is angulate posteriorly, joining the almost equally developed supra-
tympanic ridge. The crown of the head is thus a deep basin, widened
above thle tympana, and obstructed by a cross-elevation in front. Strong
ridges behind and before the orbit; nostrils vertical, a short bony longi-
tudinal ridge below them. According to tlhe characters of the genus



F :.-tiEad of Bnufo Irmsu,. (Frolm Stsjunger.)



there is no derm on tile head. Tympanumn vertically oval. Parotoids
broad oval, directed obliquely downwards. covered like the remainder of
the upper surfaces of the body and limbs, with nnmerou., closely 1
subround tubercles, with rugose surfaces. Feet rather short, wit mall
tubercles, and only one remarkably weak metatarsal tubercle, the inner.
A strong corneus ridge oi the inner margin of the tarsus. The heel
reaches the middle of the parotoidl. The toes are about half webbed.
and have a strong dermal margin. Two strong carpal tubercles. Under
surfaces studded with small tubercles, with acute points. Tongue oh-
ovate, largely free.
"The color above is a blackish-brown. the top of the head yellow
shaded: two longitudinal brown spots on the frontal region. A light,
vertebral line disappears on the back and reappears on the coccyx, and
another light line passes round the inside of the parotoids and diverges
on the scapular region. Limbs yellowish cross-banded, below dirty
white, below the vent blackish.
"This curious animal was found by George Latimer, the correspondent
of the Smithsonian Institution in Porto R.ico, W. I."
The bony crests of the head of this toad distinguish it from the intro-








SCHMI1DT, AMPHIBIANS OF I'PORTO HICO


duced marine toad at a glance, and indeed from all of the remaining tail-
less amphibians of Porto Rico.
Coloration in life*-"Tris pale brassy, sprinkled with black. General
color above dull clay-colored with a strong olive wash : blackish brown
markings and an ill-defined hourglass-shaped mark between shoulders;
also a larger blackish spot on each side of the coccyx, which is marked
by a pale streak; indications of blackish cross bands on legs; underside
dirty white, becoming flesh-colored behind and strongly reddish flesh
color on underside of femur and nearest portion of belly: tips of toes










FIG. 4.-Habitus of juvenile Bufo lemur '
(A), with side view of head (B). A. '
M. N. H. No. 10151. Natural size. ..










dark brown : tips of warts on back black, those between shoulders partic-
ularly large.
"Another specimen was colored as follows: Upper side olive, strongly
suffused with 'gallstone yellow,' which is particularly noticeable over
the insertion of the fore limbs: very few traces of dusky markings, but
the pustules are. black, especially anteriorly; an intensely orchraceous-
rufous spot on the middle of the back; on the underside the yellow suf-
fusion invades the while ground-color on the portion nearest to the
flanks.
"The third large specimen was quite similar to the last, though with-
out any rufous spot on the back, which seems to be an anomaly. Whole
upper surface darker olive, and flanks, including space at base of fore

Quoted from Sejneger, 1004. p. 72.








SCIENTIFIC SURVEY OF I'ORTO RICO


limb and below the ear, more intensely and more well-defined yellow;
underside dirty yellowish white.
"A young specimen was colored as follows: General color above drab,
more isabella-colored on head ; dark markings blackish, those on shoul-
ders pale-edged externally; flanks with a purplish suffusion and indica-
tions of a broad longitudinal band, well-defined and pale-edged above, but
gradually fading below into the pale isabella color of the belly; under-
side with a network of coarse dark-gray mottlings and marblings."
Remarks.-The five half-grown specimens (collected by myself) are
so nearly uniform and were found in so circumscribed an area that they
probably are members of a single brood. They agree in coloration with
the juvenile specimen described by Stejneger. All show the dark mark
of hour-glass shape on the shoulders. The dimensions of one of these
specimens may be compared with those of an adult recorded by Stej-
neger.
A. M. N. H. U. 8. N. M.
Parts measured No. 10151 No. 27148
Tip of snout to vent...................... .............. 37 mm. 83 mm.
'ip of snout to posterior edge of tympanumn.............. 12 12
Greatest width of head................................ 13 32
Foreleg from axilla.................... ................ 21 51
Hind leg from vent................... ................ 37 99

These five specimens were found under limestone boulders on the
artificial terrace in the moist gorge immediately behind the bath houses
of the Coamo Sprpgs Hotel. When exposed, they relied on immobility
and their resemblance to the soil for protection, and they were, in fact,
extremely difficult to see.' The first one was discovered accidentally while
I was catching a Leptodactylus under the stone beneath which it was
secreted. A large boulder sheltered three toads and one Leptodactylus.
The stomach contents of these specimens included ant remains, beetle
wings, an insect larva and segments of a small millipede. Nothing is
known of the breeding habits of this species.

Bufo marinus (Limnn)
Marine Tonl
Fig. 5

Rana marina Linnd, 175S., Syst, Nat., I, p. 211.
Bufo mariies Schneider. 1799, Hist. Amiphib., rase. I. p. 219.-Danforth, 1925,
Copeia, No. 147, p. 77.

Type locality.-America.








.9C11IJJDT, AMPIJ~~BIANS' OF P'ORTO RI CO


Disfributiotn.-Centra] America, northern South America. Native
in Trinidad, introduced in several of the Lesser Antillean islands and in
Porto Rico and Jamaica.


FIG. 5.-Bufo marinus 9. Mayagiiez. (Danforth Collection.)


During my stay in Porto Rico in 1919 I heard reports of the introduc-
tion of this species, but I did not see any specimens. Danforth's record








SCIEXTIFIC SURVEY OF PORTO RICO


from Mayagiiez seems to be the only reference in herpetological litera-
ture to the occurrence of the species on Porto Rico.
With certain reservations as to the need for revisionary studies, Bufo
marinus has an extremely wide range, extending from southern Mexico
to southern Brazil and the northern Argentine.
Diagnoss'.-A large toad with unusually large parotoid glands; bony
ridges of head well developed but low, not higher than the eyelid; no
horizontal labial ridge.
Original description.-"A frog with swollen shoulder glands and warty
hinder parts; hands with four digits, completely divided; feet with five
digits, partly divided?'
Description of a Porto Rican specimten.-(S. T. Danforth collection,
8. Mayagiiez.) Top of head bony, with well developed crests, the
height of the supraorbital crest about 2 mm. above the interorbital space,
not higher than the eyelid; orbital crests continued toward the snout by
well-defined ridges on the canthi; an anteorbital and postorbital ridge
from the supraorbital ridge about half way to the labial border; no true
labial ridge; a short supratympanic branch connects the supraorbital
ridge with the parotoid gland; no parietal branches from the orbital crests;
snout prominent, lores well-defined by the pre-orbital and canthal ridges;
eyes prominent; tympanum large, subeircular; parotoid large, distinctly
outlined, its lower edge in line with the labial border; a postrictal gland;
first finger longer than second, second as long as the fourth; toes about
one-third webbed; subarticular tubercle's single: metatarsal tubercles
small, rounded; a sharp ridge on the tarsus continuous with the outer
web of the inner toe: no tibial gland back and sides densely covered
with warts of varying size; the warts, the parotoid glands, and even the
skin between the warts are densely covered with small black spines; ven-
tral surfaces nearly smooth; upper sides of first and second fingers cov-
ered with nuptial asperities forming a fine rugose pad. Brownish gray
above; venter gray with a faint tinge of yellow, with numerous round
darker spots; throat dark gray.
A female specimen from the same locality has larger and fewer warts,
which are entirely smooth, as are the deeply pitted parotoid glands;
cranial crests somewhat lower; ventral surfaces finely granular, the
granulation finely spinose: parotoid gland more sharply outlined: fore-
limb much more slender than in the male.









SC'HMIDT, AMPHIBIANS OF 'ORTO HICO


MEASUREMENTS OF MALE AND FEMALE
S 9
Snout to vent........................................ 103 mm. i95 m.
Snont to posterior edge of tympanumn.......... ....... .11 29
Greatest width of head.................. ............. 40 37
Foreleg from axilla ........................... ......... 67 5 5
Ilindleg from vent....................... ........... 134 121
Tillia ........................................ .. .... 42 8

These two specimens are the only ones I have seen from Porto iico.
They are widely different from the Bufo maoril n of the southern half
of the South American continent. Lutz. in describing Bofo paraca emis,
suggests the composite nature of the wide-spread Bufo marines of authors.
There can he no doubt, however, that the toad here described is the one
introduced in Porto Rico, and omarins probably applies best to the ma-
rine toad of northern South America.
The marine toad is stated, by Wolcott (1921, p. 35), to have been
introduced about 1920 at Mayagiiez, by Dr. D. W. May. By 1924 it had
spread over a radius of about four miles and is apparently securely estab-
lished. A second introduction was made in 1924 at Rio Piedras, at the
suggestion of Sr. Menendez Ramos, former director of the Insular Ex-
periment Station.
Leptodactylus Fitzinger
Rana : Salo

Text Figs. 6-9

Leptodactylus albilabris (Giinther)

Cystignathn e albilabris Giinther, 1859, Ann. Mag. Nat. Hist.. (3), Vol. IV.
p. 217.-Reinhardt and Luetken, 1863, Vid. Meddel. Naturh. Foren. Copen-
hagen, 1862. p. 205.-Cope. 186S, Proc. AMad. Nat. Sci. Phila., 1868. p. 311.
LeptodactyIlux albilabris Boulenger, 1882. Cat. Batr. Sal. Brit. Mus., ). 245,
'l. 16, Fig. 4.:-Ioettger. 1892, Kat. Batr. Sanmml. Mus. Senckenherg, )p.
31.-Stejneger, 1904, Rept. U. S. Nation. Mus., 1902, p. 574, Figs. 6-14.-
Barbour., 1914, Mem. Mus. Comp. Zool., Vol. XLIV. p. 253, 255: 1915.
Proc. Biol. Soe. Wash.. Vol. XXVIII, p. 72 : 1917, idem, Vol. XXX, p. 103.-
Fowler, 1918, Papers Dept. Mar. Biol., Carnegie Inst.. Vol. XII, p. .3.
Fig. 1.-Schmidt, 1920, Ann. N. Y. Acad. Sci.. Vol. XXVIII, p). 168.-
Danforth, 1925, Copeia. No. 147, p. 77.-Noble. 1927, Ann. N. Y. Acad. Sci.,
Vol. XXX, p. 87, p. 116, Fig. 18.
Cystignathus typhonius Peters, 1876, Monatsbcer, Akad. Wiss. Berlin, p. 709.)
(not of Daudin).-Gundlach, 1881, Anales Soc. Espan. Hist. Nat., Vol. X,
p. 313.-Stahl, 1882, Fauna Puerto-Rico, p. 71, p. 161.

The writer did not hear the name ranaa" applied to this species while
in Porto Rico. "sapo" being invariably used. There is a true toad in








SCIENTIFIC SURVEY OF PORTO RI1CO


Porto Rico and, as the Leptodactylus is a decidedly frog-like animal, it is
highly unfortunate that this confusion of popular names has taken
place, since "sapo" is the Spanish equivalent of the English "toad" and
ranaa" of "frog." The word ranaa," left without definite object, is ap-
plied to various unnamed animals. At Aibonito it was used for the
ground geckos.
Type locality.-St. Thomas, Virgin Islands.
Distribution.-This species occurs almost everywhere on Porto Rico
where suitable moisture conditions exist. It is recorded from the fol-
lowing localities:

Adjuntas Catalina Plantation M ayagiiez
Aguas Buenas Coamo Springs Ponce
Aibonito Hucares Pueblo Viejo
Anasco Humacao Rio Piedras
Arecibo Lares Santurce
Arroyo Luquillo Etuado
Bayanion Mameyes El Yunque
Caguas Maricao
The loud chores of this species on the side of the Moro can be heard
distinctly from the steamer entering San Juan harbor at dawn. In
addition to the localities mentioned above, its characteristic note was
heard on Vieques Island and at Ensenada.
The record from Cnulbral appears to be the first definite one from
that island. The species occurs also in several of the Virgin Islands (St.
Thomas, St. Croix, Jost Van Dyke, Tortola, and Anegada).
Stejnegcr (1904, p. 651) regards the presence of this species as due
to accidental introduction by man. The genus was unknown from other
parts of the Greater Antilles until 1924, when Cochran described Leplo-
dactylas dominicensis from eastern Santo Domingo. Barbour (1914,
p. 253, and 1917, p. 103) dissents from Stejneger's view, chiefly on ac-
count of the number of islands included in its range. He regards the
Mexican L. labialis, which is nearly identical with this species, as an
example of chance convergence. It would be interesting to hear the
voice of the Mexican form, and to compare its life-history and larval
characters with those of the Porto Rican species.
I am unable to agree with Stejueger and Noble, who have compared
the Central American L. labialis with this species, that they are identi-
cal. In a series consisting of many hundreds of specimens from Nica-
1 A. M. N. 11. No. 10320. Culebra Island, October 5, 1919, Karl P. Sclhmidt.










SCHMIDT, AMPIHIBIANS OF PORTO RICO


ragua in The American Museum of Natural History, there is no ap-
proach to the adult size of the Porto PRican specimens: other differences
have been pointed out by Noble (1918, Bull. Amer. Mus. Nat. Hist.,


. ."


D.


p.
y.t
I'...
I~A~


FIG. G.-Leiptodatylius albilabris, to show variation in color pattern and in form of
snout. A. M. N. II. No. 10125 (A and B) ; A. 31. N. II. No. 10143 (C and D).
Natural sizp.


FIG. 7.-Leptodactylus dominiicensis. A. M. N.
H. No. 20952 for comparison with L. albi-
labrit. Three-fourths natural size.


I J -


F,


Vol. XXXVIIT, p. 323) ; and a comparison of the voices and breeding
habits of the two species remains to be made. They are so closely related,
however, that the problem of distribution is scarcely altered by this view.








NCIENTIF'IC SUiRVEY OF PORTO RICO


Leptodartyils dominiicensis is quite as closely allied to albilabris as is
the Central American form. It is certainly remarkable that this form
should have remained undiscovered in Santo Domingo until 1924.
Specimens collected.-63: Adjuntas, Aibonito, Bayamon, Caguas,
Coamo Springs, Maricao. Santurce, Utuado, El Yunque and Culebra
Island.
Diagnosis.-A smooth-skinned frog, with longitudinal glandular ridges
on the back, toes with vestigial webs, and digits tapering to a point,
without adhesive disks.
Original description.-''Tympanum distinct, one half the size of the
eye. Vomerine teeth in two short series, behind the level of the interior
nostrils. Tongue very slightly nicked posteriorly. Skin smooth, with
an indistinct longitudinal fold on each side; a transverse fold between
the fore-legs, another across the posterior third of the belly. Snout
moderately produced. Tarsus was a longitudinal fold; interarticular
tubercles prominent. Male with two vocal sacs, communicating with
each other, each with a separate slit. A white or whitish streak round
the snout to the axil.
"Colour of the adult:-Above uniform dark bluish-black; the upper
leg with some black cross bars superiorly, and some whitish spots
posteriorly. The lower parts white, the throat speckled with brown.
The labial streak whitish, indistinct below the eye.
"Colour of the young:-Brownish olive marbled with darker; uniform
white inferiorly; the labial streak white, very distinct.
"These descriptions of the colours are taken from quite fresh speci-
mens in spirits.
"Hab. St. Thomas. The specimens are now in the British Museum."
There is no question of the specific identity of the St. Thomas
Leptodactylus with that on Porto Pico. To Guinther's description may
be added the following-the vomerine teeth are in curved; slightly oblique
series; nostrils nearer the tip of the snout than the eye; tympanun about
two-thirds the diameter of the eye; first finger much longer than second,
and second and fourth fingers equal; toes slightly webbed at base; two
metatarsal tubercles, the inner connected with the slight tarsal fold;
heels overlapping when the limbs are placed at right angles to the body.
Coloration of a living specimen.*-"General color above olive, the
dusky markings dark grayish brown, nearly blackish brown below the
dorso-lateral fold and on femur; the dorso-lateral fold and narrow
edges around the dark markings pale olive gray: the transocular band
Quoted from Stejneger, 1I-04, p. 576.








SCN'HllD/T, AMPI'IIHIlA OF /'PORTO HIICO


and cutting edge of lip dark grayish brown; the supra-labial light band
pale straw yellow: underside whitish; throat finely sprinkled with dark
chocolate brown; iris olive silvery, overlaid with blackish."
The form of the snout exhibits a striking variation, some specinlns
having a decidedly flattened and sharp-sided snout. The possibility that
this snout form is correlated with a tendency to burrow more or less
offers an interesting problem for observation in the field.
This species exhibits a great variability in coloration with a relative
uniformity in structural characters. Fowler (1918, p. 3. Fig. 1) has
figured the extremes of color pattern in Porto Rican specimens. Seven
of the fifty specimens of the present series have the broad median stripe
on the back, the others varying chiefly in the distinctness of the dorsal
Y-shaped markings. The measurements of the largest specimen and of
one of apparently recent transformation are as follows:
A. M. N. H. A. M. N. H.
Parts measured No. 10182 No. 10034
Tip of snout to vent.................................. 49 mm.f 16 mm.
Tip of snout to posterior edge of tympanum. ......... 18 7.5
Greatest width of head ................ .............. 17 7
Foreleg from axilla................................ 29 10
IIind leg from vent to tip of longest toe.............. 78 24

Leptodaclylus albilabris is partial to moist situations, from sea level
to an altitude of at least two thousand feet. In towns and villages it is
found in the drains and gutters, regardless of filth, wherever a little
muddy water accumulates. It is especially abundant in moist meadows
or in irrigated cane fields. In the coffee plantations, the adults are
found concealed beneath logs or stones, while the young hop about
everywhere on the ground, apparently feeding.
Of 25 stomachs examined 8 were empty. Of the remainder. 4 con-
tained land snails; 2 contained spiders (1 a large Lycosid spider and egg
sac) ; 2 contained ants; 2 contained beetles; 2 contained bugs; 2 con-
tained flies (Muscidae) ; 1 a small moth; 1 a large caterpillar: 1 a me-
dium-sized cockroach, and 7 unidentifiable insect remains.
The nest of this species was observed by Stejneger (1904, p. 579)
under a flat stone in a stream. Peters (1817, Monatsber. Akad. Wiss.
Berlin, 1876, p. 709) records one observed by Gundlach in a "wet bur-
row." At Coamo Springs, on the terrace behind the bathhouses of the
hotel, the water of some of the springs forms a permanent rivulet at the
base of the cliff. Leptodactylus albilabris was abundant on the terrace,
beneath loose stones, and under a large stone at the edge of the creek.
t 144 mm., given by Stejneger (1904, pp. 570, 578) is obviously a mnisprint.








SCIENTIFIC' SURVEY OF PORTO RICO


the writer found, on August 27, 1919, a shallow, rounded excavation,
G or cm. in diameter and about 3 cm. deep, filled with a mass of white
foam, in which were the small tadpoles of this species (12 mm. in length,
body 3-4 min.). There were between 75 and 100 tadpoles in the mass,
by no means confined to the central hollow, which was present, as in the
foam-mass described by Stejueger. The bottom of the excavation was
about 3 cm. above the water level. Two similar excavations were dis-
covered in the immediate vicinity, in the same relative position with
reference to the water, but empty. On August 29, near Bayamon, a small
mass of foam. between 3 and 4 cm. in diameter, containing similar tad-
poles was noted under a stone on a hilltop, with no water whatever in
the neighborhood. On October 1, near the Forester's Cabin on El
Yunque, at about 1200 feet, a nest of this species was observed beside a
pool of standing water (also at a slightly higher level than that of the






FIG. S.-Lateral view of tadpole of Leptodluctylus albilabris. (After Noble.)

water) under a rotten log. This nest contained between 150 and 200
eggs, uniformly distributed through the foam, and with no central hol-
low. It was somewhat larger than those previously observed, measuring
8 cm. in diameter. The eggs are light yellow, and measure 2.5 mm. to
3 mm. in diameter. The smallest tadpoles taken swimming at large
measured 6 mm. in body length, which probably represents their size at
the time they escape from the foam. It is evident that the tadpoles
usually will be washed from the nest into the adjacent water by a flood or
heavy rain. The location of the small nest away from water was prob-
ably a mistake on the part of the frog, and the nest described by
Stejneger under water probably had been covered by a rise in the creek
after the deposition of the eggs. The largest larvae, nearly ready to
transform, measure 13 mm. from snout to vent. The V-shaped dorsal
markings are already evident in the tadpoles at this stage. The median
dorsal white line is probably an adult character.
Description of tadpole.*-"Length of body about once and one-third
its width and slightly less than one-half the length of the tail; nostrils
nearer the eyes than the end of of the snout: distance between eyes one-
fifth more than distance between the nostrils, and considerably less than
Quoted from Stejneger, 19114, p. 577.








CHMIIDT. AMII'HIBIA A' OF PORTO RICO


width of mouth : distance between nostrils equals their distance from
eyes, as well as the diameter of the eyes: spiraculium on left side. directed
backward and upward, situated above a line drawn between the base of the
munseular part of the tail and the mouth, and nearer to the posterior ex-
tremity of the body, being about halfway between anterior border of
eye and insertion of hind legs: anus a long tube. median and larger
than the spiraculum: tail about four times as long as deep, ending in
an obtuse point: both upper and lower crests confined to the tail and
nearly equal in depth, their edges being nearly parallel until the ter-
minal third ; the depth of the muscular part of the tail at its base about
two-thirds the greatest total depth."
The chorus of this species is one of the most insistent notes heard in
Porto Rico, both by day and night. though somewhat stronger by
night. Stejneger compares the note to a rapidly repeated "pink-pink-
pink"; .it is comparable in quality with that of the North American




FmI. 9.-Mouth parts of tadpole of Lep- 0)
todactylas albilabris. (After Stejneger.) .......,





Acris. The interval between notes is variable, but usually very short
and regular. There is an occasional somewhat guttural trill. The note
ceases on the near approach of a person, and it is exceedingly difficult to
discover the singing individual, for the creature may be concealed be-
neath the edge of a rock or stick, or be flat on the ground between the
roots of the vegetation.

Eleutherodactylus Fit zinger

The present state of our knowledge of the Porto Rican species of
Eleutherodactylus does not justify the attempt to draw up a synopsis
of the species in key form. Four of the ten species are imperfectly
known,-uiicolor and locustus based on single specimens, briitoni and
cramploni on small series,-and the most useful of all characteristics in
life, the voice, is known for only six of the species, the breeding habits
for only a single species! The difficulty in distinguishing species of this
genus does not, however, vanish completely, even with the accumulation
of large series. Eleutherodactylus is plainly one of the groups in which








,,I('i IT'/FC NURt'EYl OF PORTO ICl('O


speciation is notably active, and like Sceloporu.i among the lizards, its
study may be recommended "as an excellent pire de resistance for those
persons who do not believe in the doctrine of derivation of species."
The genus falls into two groups in Porto Rico,-on the one hand E.
riclimondi and E. moniiesis, having long transverse series of vomerine
teeth, and on the other, species in which these series of teeth are short
and oblique. In the latter group, E. unicolor occupies an isolated posi-
tion by reason of its posteriorly placed nostrils and widely separated
vomerine teeth.

Eleutherodactylus portoricensis Svhmiidt
Coquf
Text Figs. 10-13

Hylodes martini.(eci.i.8 Peters. 1876. Monatsber. Akal. Wiss. Berlin, 11. 709.,
1'1. I, Figs. 1-5, 7-9 (not of Tschudi).-Gundlach. 1881, Anales Soc. Espail.
Hist. Nat., Vol. X, p. 315.-Stahl, 1882. Fauna Puerto-Rico, p. 71. p. 101.-
Garman, 1887, Bull. Essex Inst., Vol. XIX. p. 13.-Boettger, 18,02, Kat.
Batr. Mus. Senckenberg., p. 29.
Eleuthcrodactylus auriculatus Stejneger, 3904. Rep. I. S. Nation. Mus.. 1902.
p. 583. Figs. 15-19 (not of Cope).-Fowler. 1918. Carnegie Inst. Wash.
Publ. 252, p. 4, Fig. 2.-Schmidt, 1920, Ann. N. Y. Acad. Sci., Vol. XXVIII.
p. 170, Figs. 1-2.-Danforth, 1925, Copeia, No. 147. p. 77.
Eleutherodactylus portoricensis Schmidt, 1927. Amer. Mus. Novit., No. 279, p. 2.

The native name, coqui, expresses the repeated element in the call of
this species very satisfactorily.
Type locality.-El Yunque, 2000 feet altitude.
Distribution.-This is the common Porto 1'iean tree frog, recorded
from very many localities. The places where it occurs, compiled from
my list and Stejneger's records, are as follows:

Adjuntas Coamo Springs Ponce
Aguas Buenas hiumacao Rio Piedras
Aibonito .1 ayuya Santurce
Alto Mauzano Lares Utuado
Areciho Luquillo Vega Baja
Caguas Marmeyes EHI Yunque
Catalina Plantation Maricao
Catafio Mayagiiez

There seems to be no record from southwestern Porto Rico.
Specimens collected.-207: Adjuntas, Aibonito. Alto Manzano, Catafio.








SCHMIDT, AllI'HIBIANS OF PORTO HIGO


Coamo Springs, Jayuya. Maricao, Rio Piedras, Santurce, Vega Baja and
El Yunque.
Diagnosis.-An Eleatherodactyls of moderate size and stocky habitus;
vomerine teeth in two short oblique series, behind the ehoanas; nostrils
near the tip of the snout: tympanum about half the diameter of the eye;
disks of toes about equal to those of fingers, about three times as broad
as the narrowest part of the corresponding phalanges; no trace of web;
ventral disk faintly indicated; tibia less than half the length from snout
to anus; concealed parts of thighs immaculate, reddish in life.
Original description.-"Ilead broader than body, its width slightly
greater than the distance from tip of snout to the posterior border of the
tympanum; nostril twice more distant from the eye than from tip to
snout: diameter of eye equal to its distance from the nostril; tympanum
nearly half the diameter of the eye. a little broader than its distance from



FIG. 10.-Inside of mouth 6
of Eleutherodlactylus por-
toricensis (left) and of
E. richimondi (right),
showing different ar-
rangement of the vomer-
ine teeth. (After Stej-
neger.)




the eye; heels broadly overlapping; heel reaching the eye when the limb
is laid forward along the body; canthus rostralis rounded but well de-
fined; lores sloping, slightly concave; disks of fingers subequal; disks of
toes subequal, very little smaller than those of the fingers; first and second
fingers equal in length : first and second toes subequal; skin finely rugose
above, with a narrow median raised line, the general effect smooth; belly
and thigh granulate; a fold across the chest; ventral disk faintly indi-
cated by an impressed lateral line; vomerine teeth in straight, short,
oblique series, their distance from the choanse, in line with their outer
borders, a little less than their length, the distance between them about
half the length of one series. -t-
"Color dark brownish gray : a light eanthal line over the edge of the
eyelid, broadening over the tympanum into a dorsolateral light band,
which merges into the light belly color posteriorly; concealed surfaces of
thighs and tibi? immaculate (reddish in life)."










46 SCIENTIFIC SURVEY OF PORTO RICO

MEASUREMENTS OF TYPE
Length of Body...................... ........... ............... 36 m m .
W idth of H ead ............................... .. ................. 15
Tip of Snout to Posterior Border of Tympanum..................... 14
Length of Arm ............... .............. ............... 21
Length of Leg.................................................... 52
Tibia .............................. ........................... 17
Tym panum ....................................... .............. 2
Eye ................................. .............. ........... 4.5
Largest Finger D isk.............................. ............... 2


FIG. 11.-Three common color variants of Eleathcrodartylts portoricensts. A, A. M.
N. II. No. 10139; B, No. 10243; and C, No. 101240. Natural size.








St'CHIIDT, AMPHIBIANS OF PORTO RICO


Remaurks.-The only noteworthy discrepancy between my description,
quoted above, and the detailed description of a Porto Rican specimen by
Stejneger (1904, p. 585) is in the size of the disks of the toes, which I
find to be nearly as large as those of the fingers, but which are stated by
Stejneger to be smaller, and are figured by him as only about half tl.e
size of the finger disks. I am convinced that this difference rests on a
different preservation of material or possibly on actual variation. Fow-
ler's figures exhibit a good deal of variation in this respect. The ventral
disk in the specimen described is very faintly marked, and the difference
in this character is probably merely one of opinion.
Coloration in life."'-"Above dusky fawn color with a very narrow
vertebral line, a narrow canthal line and a broad lateral line in continua-
tion of the latter, pale buffy pink, all the light lines and bands more or
less edged with dusky; triangle on top of head forward of a line through
the center of the eyes distinctly paler, a dusky cross-line at middle of
eyes defining the triangular space behind; below pale, greenish on belly;
underside of femur dull ferruginous: iris golden, shaded with reddish
and brownish and reticulated with blackish."
A specimen of Eleuthierodactylus auricul tus from El Periu. Monte
Libano, Guantanamo, Cuba, loaned for examination by Dr. Thomas
Barbour, is much smoother above than E. portoricensis, notably on the
eyelids and snout.
E. portoricensis is remarkable for its color variation, with a compara-
tively stable structure, but a considerable variation in measurements. A
specimen in the collection of Professor (4. E. .Tohnson of the University
of Porto Rico, collected by him in the Luquillo Forest, is noteworthy for
its size, being apparently a giant individual. The maximum size in more
than three hundred specimens in the National Museum and the present
collection is 44 mm. from snout to vent, while Professor .Johnson's speci-
men measures 52 mm. Its measurements compared with the largest in
the present series are as follows:
A. M. N. H. Johnson
Parts measured No. 10241 specimen
Tip of snout to vent..................................... 44 mm. 52 nim.
Tip of snout to posterior edge of tympanum................ 18 20
Greatest breadth of head................... ............. 19 23
Foreleg from axilla...................... .............. 29 35
Hind leg from vent.................... ............. .... 69 80

The most frequent coloration is a grayish brown of varying shade,
sometimes reddish, sometimes nearly black. This may be uniform or

Quoted from Stejneger, 1904, p. 585.








SC)NT'IFIC KSUURVEY OF PORTO RICO


mottled with darker. In the lighter specimens there is nearly always
a dark interorbital mark. and in a few the snout is white in front of this
either with a broad transverse white band or completely light to the tip
of the snout. There is usually also a dark sub-canthal mark, interrupted
by the eye, and continued over the ear for a short distance. In a few
cases the dorsum is spotted irregularly with vivid white spots. In 18
out of 194 specimens a light line, beginning at the snout and passing
over the edge of the eyelid to the ear, continues as a broader light dorso-
lateral band to the thigh. In 19 specimens there is a sharp median white
stripe (compare Fowler, 1918, Fig. 2). In 5 there is a broad median
light band, about four times as broad as the more common narrow line.
The hind legs are occasionally distinctly barred, more usually indistinctly
barred or uniform. The concealed surfaces of the thighs are often





/\ VIG. 12.-Embryo of Eleutherodaetylus portori-
censis. A. M. N. H. No. 10302. Six times
S/Inatural size.






bright pink or red. The venter is usually light and unspotted, occasion-
ally it is spotted with groups of dark-brown punctations. In no specimen
were the concealed surfaces of the thighs reticulated with the fine or
coarse dark network of E. antillensis. The color variation has also been
noted by Stejneger, but I am sure that the juvenile variant described by
him from El Yunque must refer to the dwarf species, E. gryllus.
The majority of the specimens collected were taken at night, either
singing or merely sitting about on vegetation, sometimes several feet
from the ground. Others were found concealed under logs and stones,
especially in coffee plantations. The most usual hiding place of this
species in the daytime is beneath the sheaths of the outer leaves of the
banana. Specimens in this situation are almost without exception
uniformly colored and nearly black. About every third banana plant
examined was inhabited by one or more of these tree frogs.
Persistent search about the banana plants failed to discover the eggs
of this species, and it was not until the writer visited the Luquillo








SCHMIDT7, AMIJPHIBIANS OF PORTO RICO


*Forest that a single egg-mnass was discovered in a basal leaf if an air
plant, just at the surface of the water in the lower part of the leaf. A
large E. auricilatus in the same plant, but not on the same leaf as the
ego-s, escaped. There were thirty-six eggs. with well-advanced embryos,
adhering in an oval mass from which individual eggs could easily he
detached. The eggs measure 6-8 mm. in greater diameter, being some-
what elongated in the axis of the embryo.
The young of this species are extraordinarily abundant, and it is
difficult to understand why the eggs are so infrequently observed. It is
possible that at the time of the writer's visit (August-October) the
height of the breeding season was past. The only date of breeding previ-
ously recorded is that noted by Gundlach, May 24 (Peters, 1877, Monats-
ber. Akad. Wiss. Berlin, 1876, p. 709). Professor Johnson found a mass
of eggs, with embryos at about the stage of those observed by the writer,
in the same bunch of moss in which the giant female specimen, men-









FIG. 13.-Peter's figures of the embryo of ElIctherowdutylus portoriceltuii.
tioned above, was collected, July 8. 1919. Gundlach (loc. cit) also ob-
served a female sitting on the egg-mass received by him, while Bello y
Espinosa (Martens, 1871, Zool. Garton, Vol. XII, p. 351) records that in
the case noted by him the parent frog remained in the neighborhood of
the eggs "as if to guard them." From these several observations it
appears not unlikely that the female does remain in the neighborhood of
the eggs until they are hatched. but further observations on this point
are desirable. Ruthven (1915, Occ. Papers MIns. Zool. Univ. Michigan,
No. 11), observing the breeding habits of E. cruenlus Peters, in Colombia,
found no evidence of such a habit.
The chorus of this species, or the isolated notes (f single males, are
among the most familiar and insistent sounds in Porto Iico. The call
is a clear, whistled co-qui. co-qui, varied by co-qui-quf-qui-qui-qui.
Two males often call alternately from neighboring stations. A favorite
situation for the singing male is the base of the leaf of a lilliaceous
plant or the center of a whorl of leaves on shrubs, etc., which appears to
magnify the sound and thus increase its ventriloqnial character.








SCIEXT'IFIC l$URVEY OF PORTO RICO


For its relations with other West Indian Elettherodactylvs. and the
history of the species. I may quote my remarks when proposing a name
for this species:
"It is paradoxical to write of the most abundant species of tree-frog
of Porto Rico as new to science, for this form is one of the best known in
the genus, represented in many museums by large series of specimens.
It is really famous, for its eggs and embryos were the basis for the article
by Peters, describing the direct development, with suppression of the
tadpole stage, which is a general character of the genus Eleuthlerodac-
tyius. Peter's figures have found their way into great numbers of text-
books, usually under the original designation, fTylodes ma.rtiicetnis.
"The name now proposed for this well-known and well-characterized
species is, nevertheless, the first to be based on Porto Hican specimens.
The confusion of this form with other Antillean species has been due
to the weight of authority that has identified it first with the Lesser
Antillean Eleulherodactylus martinicenii.s (Tsehudi)-Peters, Gundlach,
Garman, and Boettger-and, later, with the Cuban Eleutherodactylus
auriculatus (Cope)-Boulenger, Stejneger, and Barbour. In dealing
with this species in 1920, I accepted the identification with auriculattus
without question.
"Stejneger, in 1902, accepted Boulenger's record of auriculatus from
Santo Domingo, and its occurrence on that island would of course make
its presence in Porto Rico much more probable. The Nobles secured
an allied species in the Dominican Republic, described by Dr. Noble as
Eleutherodactylus auriculatoides (1923, Amer. Mus. Novitates, No. 61,
p. 3) and it is probable that this species represents auriculatus in Santo
Domingo.
"The renewed and more intensive study of the Greater Antillean
amphibian faunam was, to some extent, initiated by my field work in
Porto Rico in 1919, which added six species of Eleatlerodactylus to the
supposedly well-known herpetological fauna of that island. This was
followed by the work of Dr. and Mrs. G. K. Noble in the Dominican
Republic in 1922, which added five new Eleut/ierodaclylus and a new
Hyla to the IIispaniolan fauna. The recent additions to the Cuban tree-
frogs (eight species) and to the Jamaican fauna (six Eleutlerodactylus
and a Hyla) by the field work of Dr. Emmett R. Dunn in 192 1 and 1925
were, consequently, scarcely surprising, though it may be emphasized that
all of these islands were supposed to be well explored herpetologically.
The new crop of novel species was due to the application of a simple
technique of collecting by voice at night, using an electric flashlight.









,'<'Hi.fllT', A.i11'P1//.iAN OF I'PORTO HI('O


"A better knowledge of the old species has inevitably accompanied
the recognition of the new forms., and it is nuo evident that there are no
native species of this genus generally distributed in the Greater Antilles.
The Cuban EleutherodnctIlu.s ouriculatus is now well known through Dr.
Dunn's field work. He writes me that this species does NOT breed in
bromeliads. and( that its cry resembles the syllables "chi-leen." The
repeated "'eoqui" of the Porto R]ican species, which gives it its native
name, is one of the most characteristic sounds of the nocturnal chorus
in Porto Rico.
"All of this contributes little liv little to the certainty that the common
Porto Rican tree-frog is specifically distinct from any other West Indian
formn"
Eleutlierodaetylus gryllus S-.himidt
Text Fig. 14
Irutithecroetiuctlu, cry'lus Schmidt. 192(0. Ann. N. Y. Aeal. Sci., Vol. XXVIII,
p. 172. Fig. 3.

There is no native name for this species.
Type locality.-El Yunque, near the Forester's Cabin, about 1300 feet
altitude.






FIG. 14.-Eleutherodactylus gryllus. A. M1
N. II. No. 10226. Twice natural size. c






Di.tribution.-Eleiutherodaciylus gryllus is confined to Porto Rico,
where it is known from the Luqnillo Forest and from Maricao. It is
probably confined to the more humid higher parts of the island.
Specimens collected.-16: Maricao and El Yunque.
Diagnosis.-Distinguished from Eleu.therodacdllus porloricensis by its
shorter snout, less granulate venter, and its minute size.
Description of type.-"Habitus of Eleitlerodaclylus portoricensis, but
with a distinctly shorter snout, its length equal to the diameter of the
eye (in E. portoricensis the diameter of the eve equals its distance from








SCI'ETIFIC SUI'VEY OF I'ORTO RICO


the nostril), and to the interorbital space; canthus rostralis rounded;
nostril one-third the distance from tip of snout to eye; tympanum
scarcely distinct, one-fourth the diameter of the eye. its distance from the
eye equal to its diameter; toes without vestige of web; digital disks well-
developed; first toe as long as the second; an inner and outer metatarsal
fold; vomerine teeth in two oblique patches behind and within the
choanae: tongue large, slightly nicked behind; skin smooth above, but
apparently much more glandular than in E. portoricensis; venter
strongly granulate; a large subgular vocal sac.
"Middle of the back, beginning with an interorbital line, dark gray,
enclosing a light spot on the occiput: sides and snout lighter, the darker
color everywhere consisting of minute black punctations, especially evi-
dent on the limbs and throat; venter light."
MEASUREMENTS
Tip of snout to vent ................................ .............. 16 nun.
Tip of snout to posterior border to tympanum......................... 5.5
greatest t breadth of head................................... 6
Foreleg from axilla .............................. ............... 11
Hind leg from vent............................................... 24
Tibia ...................... .................... ............... 8.5

Renmarks.-The type is a male, taken singing at night, with the
usual pale night coloration. Specimens taken in the daytime (concealed
under moss) are very dark in color and exhibit considerable variation in
pattern, two having a light median dorsal line. In a specimen taken in an
air plant (Xo. 10291) the dorsal dark area is cinnamon brown and the
sides bright pale green, the legs with dark bars; this coloration has been
described by Stejneger (1904, Rept. U. S. Nat. Mus., 1902, p. 586) as
a variant coloration of juvenile E. auriculatus ( portoricensis). The
darker specimens have narrow light crossbands on the limbs. The
granulation of the venter in the female specimens is faint, though still
evident.
This species was very numerous at Maricao and on El Yunque, sing-
ing frequently from trees, at least ten feet from the ground. On El
Yunque specimens were collected in air plants, near the peak. and under
moss on the rocks of the peak itself.
The song is a rapid succession of shrill clicks, very insect-like, the
chorus sounding not unlike the rapid clicking of a telegraphic instru-
ment.
Were it not for the minute size of the singers, and the extremely dis-
tinct note, this species might well be considered the young of E. portori-








'JID1T11)7'. AMI'llHIBANS OF PORTO'0 RICO


cpiissl; I ain unable to agree with Stejnegerls supposition that its note is
made by juvenile males of the latter species. The gonads. at any rate.
appear to be those of an adult in the specimens examined. differing in
form and pigmentation from those of young E. portoricedi.is of similar
size.
Nothing has been added to the knowledge of this species since its de-
scription in 1920.
Eli tlilerodl rtyluis locuslns Siliniilt
Text Fig. 15
Elruthctrolductulus locusrin.s Slihmlidt, 1920, Ann. N. Y. A.nl. Sci., Vol XXVIII,
p. 174, Fig. 4.

No native name is known for this species.
Type localily.-El Yunque, near the Forester's calin, about l:Ii0l) feet
altitude, Tjnquillo Forest Reserve, Porto Rico.
Distribution.-Known only from the type locality.
Specimens collected.-1 : El Yunque.







FIG. 15.-Elclitheroductylus locu'stus. A. A1.
N. HI. No. 10240. Twice natural size.







Diagnosis.-Size small: snout obtuse: nostril much nearer to the end
of the snout than to the eye: tympanum small, indistinct, one-fourth
the diameter of the eye. separated from the eye by a little more than its
diameter; vomerine teeth in two oblique series, behind and within the
choanme; toes free; digital disks well-developed: tibiotarsal articulation
reaching the posterior border of the eye; heels overlapping when the legs
are placed at right angles to the body; skin rugose above, with scattered
round tubercles, especially on the eyelid; venter smooth ; inner face of
thighs finely rugose.
Original description.-"Head slightly longer than broad, slightly nar-








N'IE'VT''IFIC SR'IVEY OF PORTO RICO


rower than the bod : snout moderately obtuse, its length anterior to the
eye exceeding the interorbital space; nostrils one-fourth the distance be-
tween eye and tip of snout from the latter; tympanuim scarcely distinct.
one-fourth the diameter of the eve. separated from the eye lby a little
more than its diameter: eanthus rostralis rounded elbow and knee
pressed along the side. overlap; heels overlap when the legs are placed
vertically to the axis of the body; tibiotarsal articulation reaching the
posterior order of the eye ; disks of fingers and toes well-developed; toes
without vestige of wel): inner and outer metatarsal tubercles present: no
tarsal fohl: fir.t toe as long as the second : vomerine teeth in two linear
oblique patches, converging posteriorly, well-separated on the median
line, behind and within the choanmi by about the diameter (it the choana:
tongue large, slightly nicked behind : skin rugose above, with rounded
tubercles: a well-marked mid-dorsal ridge from snout to vent; eyelid
strongly rugose: venter smooth (faintly rugose under the lens) thighs
slightly rugose; male with a large subgular vocal sac.
"Dorsum gray mottled with grayish brown : a well-defined inter-
orbital dark bamnd: sides of canthus with a dark mark, interrupted by the
eye, extending over the tympanum; legs not barred, with dusky mark-
ings: venter uniform, light."
Al EASi'REMEINTS
Tip of snout to vent..................... ...... ............. 19 llm.
Tip of snout to posterior edge of tyipiiminum....................... 7
Greatest breadth of head ............ ........................ G.
Foreleg from axilla............... ........................ 12
Hind leg from vent ............................... ............. 20
Tibia ................................................ ................ 9

Remud's'.-This species was discovered hv accident, singing on a leaf
some three feet from the ground. Its song is the most distinctive of any
noted in Porto Rico, beginning with a shrill continuous note almost at
the limit of nudibility, which is followed by a succession of clicks. So
closely does this note resemble a familiar type of note produced by long-
horned grasshoppers that the writer neglected to search for the author
of the sound, and watched the present specimen repeat the song several
times before being convinced that it really proceeded from an Eleath ero-
d(actylus.
Eleullierodaclylus locustus is closely related to E. porloricensis, from
which it is distinguished by its small size and smooth venter. Even
more closely related to tlie still smaller new species E. gryllus. it is
nevertheless readily distinguished by its smooth venter and more rugose
dorsum, as well as by its remarkable voice.








SC(IIMIDT, AI'HIBIAXNS OF PORT RICO


This species has not been rediscovered since its description in 1920.
I might entertain a doubt as to its validity, in view of its being based
on a single specimen, had not Dr. and Mrs. Noble since observed an
Eleuthlerodactylis with a similarly peculiar note in Hispaniola.

Eleutliherodaetylus cramnptoni* Sclmidt
Text Fig. 16
Elcutheroductylis crrmptoni Schmidt. 19220, Ann. N. Y. Acad. Sci.. Vol.
XXVIII, p. 17i;, Fig. 5.

No native name is available for this species.
Type locality.-Peak of El Ynnque, 3485 feet altitude, Luquillo
Forest lReserve, Porto Rico.
Distribution.-Known only from the type locality.
Specimn Ps collected.-3: El Yunque.







FIG. 10.-Eleulherodactylis cramptoni. A. ..Y I
M. N. II. No. 10305. Twice natural
size. '" "







Diaonosis.-Size small: habits stout; hind legs short: snout very
obtuse, canthus rostralis rounded; dorsuim very rugose with rounded
tubercles; vomerine teeth in two oblique linear series, extending laterally
as far as the inner border to the choanw: digital disks large; uniform
dark brown above, light brown beneath.
Oriyinol descripion.--"Habitus stout, compact: snout short, obtuse.
canthus rostralis rounded: nostril one-third the distance from tip of
snout to eye; heel reaching the anterior border of the orbit; heels meet
but do not overlap when the legs are placed at right angles to the body:
both anterior and posterior limbs notably stont, nearly twice as thick as

Named for Professor Henry E. Crampton, whose active interest and investigation
have greatly furthered the zoological work of the Scientific Survey of Porto Rico and
the Virgin Islands.








SCIENT''IFIC SURVEY OF PORTO RICO


those of E. portoricensis of the same body length; vonierine teeth in two
linear, oblique series, extending laterally as far as the choane; tympanum
small, distinct; dorsum covered with rounded tubercles, extending onto
the eyelids and snout; venter finely granular; digital disks large, first
toe as long as the second; no subgular vocal sac.
"Color uniform brown above, lighter brown below, slightly variegated
with lighter punctation."
)IEASUREM E.NTS
Tip of snout to vent........................... ............... .. 19 mm.
Tip of snout to posterior edge of tympanum....................... 6.5
Greatest breadth of head.................................. 7
Foreleg from axilla................ ...... .... .............. 12
Hind leg from vent ................ ....................... 29)
Tibia ............................ ............................... 9

Remarks.-The two paratypes are similar in every respect to the type,
with the single exception that one of them is slightly more mottled with
light, and has the hind legs indistinctly barred.
All three specimens were taken under moss in the crevices of the rocks
on the peak of El Yunque.
The species is a well-differentiated one, characterized by the stoutness
of its limbs, the obtuseness of the snout and the extreme rugosity of the
dorsum.
Like the other novelties in my collection of 1919, this species awaits
the verification of further field-study.

Eleutherodactylus antillensis (Reinhardt & Luetken)
Coqui
Text Fig. 17
Hylodcs (ntillensis Reinhardt and Luetken, 1863, Vidensk. Mled. nature. For.
KjSbenhavn, 1862, p. 209.
Elcutherodactylus antillensin Stejueger, 1904, Rept. U. S. Nat. Mus., 1902, p.
591, Figs. 20-24.-Barbour, 1914, Mem. Mus. Comp. Zool., Vol. XLIV, p.
247;: 1917, Proc. Biol. Soc. Wash., Vol. XXX. p. 102.-Schmidt, 1920, Ann.
N. Y. Acad. Sci., Vol. XXVIII, p. 178, Fig. 6.
HIylodes nmartinicnsi.v Peters, 1876, Monatsber. Akad. Wiss. Berlin. 1576. 1'1 .1.
Fig. 6.

This species is not distinguished by the Porto Ricans from the E. por-
Ioricensis, and when observed at all is called a "coqui."
Type localily.-St. Thomas, Virgin Islands.
Disl.rib ion.-This species was long known only from the Virgin
Islands, St. Thomas and Tortola, and from Vieqnes Island. It is








SCHMIDT, AMPIHIHIANS OF PORTO 1?]CO


doubtfully reported from St. John and St. Croix. I collected it on Porto
RIico and Culebra in 1919.
Specimens collected.-30: Aibonito, Bayamon, Alaricao, Santurce and
('ulebra Island.
Diagnosis.-Limbs shorter than in Eleulherodactylua portoricensis,
the heels just meeting when the legs are bent at right angles to the
body. Posterior surfaces of the thighs with a dark reticulation. Venter
more coarsely granulated and digital disks smaller than in porloricensis.
Original description.-"An Hylodes with a verrucose venter, palatine


Fit. 17.-Eleautherodactylus antil-
lcnsis. A. M. N. H. No. 10010.
Twice natural size.


teeth moderately separated, each series wedge-shaped, the two forming an
angle open forward; [palatine teeth] widely separated from the border
of the lip; digital disks rather large."
Remarks.-Stejneger* gives a detailed description of this species, based
on a Vieques Island specimen:
"Tongue rather broad, heart-shaped, slightly nicked behind: vomerine
teeth in two club-shaped oblique series, some distance behind but not
laterally beyond the choanie, converging backward and well separated;
nostril much nearer the tip of snout than the eyes, their distance from the
eye less than the diameter of the latter: upper eyelids narrower than
the interorbital space; tympanum a little less than one-half the diam-
eter of the eye, its distance from the eye less than one-half its diam-
SStejneger, 1904, p. 592. 71J .

*.. t;
I Z "' '








S('IE7NTIFIC URVEY OF PORTO RICO


eter; fingers with rather small disks, first equalling second ; disks of
toes not smaller than those of the fingers; tip of first toe reaching disk
of second; two metatarsal tubercles, the outer being rather small and
obscure: series of plantar tubereles corresponding to metatarsals; no
tarsal fold: the bent limbs being pressed along the side, knee and
elbow, fail to meet; hind limb being extended along the side, heel
reaches the eye; hind limbs being placed vertically to the axis of the
body, the heels barely meet; skin above with scattered granules and a
very narrow raised median line from tip of snout to vent; throat and
chest smooth, belly and posterior aspect of femur strongly granular:
a strong fold across the breast between the axilhl."
The original description is sufficiently diagnostic as the only other
species on St. Thomias is the smooth-skinned E. lentus, which has the
vomerine teeth in long transverse series.
Color.-In coloration this species is less variable than E. portoricensis
but the median white dorsal line may be present or absent. It is de-
veloped in twelve of the present specimens. The usual color is grayish
brown, with faint dusky markings, and a sharply defined black canthal
line which extends over the ear and a short distance beyond it, outlined
above in most eases by a very narrow white line on the canthus extending
over the eyelid. The concealed surfaces of the legs are reticulated with
black, which affords a fairly good character for distinguishing this species
in the field from E. portoricensis. One specimen, No. 10001, a male, is
violet-red above, has a very heavy black canthal and supra-auricular
mark, and the concealed surfaces of the legs black with sharply defined
white spots.
The measurements of the largest specimens of each sex follow:
Parts measured No. 10117 dc 10082 9
Tip of snout to vent.................................... 24 mm. 33 mm.
Tip of snout to posterior edge of tympanum .............. 10 13
Greatest breadth of head................................ 11 14
Foreleg from axilla...................... ............... 16 19
Hind leg from vent................. ................... 38 48

Habits.-This species prefers slightly lower herbage and' slightly wet-
ter situations than E. portoricensis, which is often associated with it. It
ranges to an altitude of nearly 2000 feet at Aibonito.
Nothing is known of the breeding habits of this species.
The song of E. antillensis may be readily distinguished from that of
E. porlori censis by its more metallic quality, and the frequent series of
uniform notes ki-ki-ki-ki-ki - -. The males conceal themselves more








SCHIllDT. AMPHIBIANS OF PORTO RICO


carefully when calling, making it exasperatingly difficult to locate the
singer. They often sing from a position in the axils of the leaves of
lilliaceous plants. In Santurce along the railroad and trolley embank-
ments north of the Hotel Eureka, I found this species more abundant
than E. portoricensis. The single specimen from Culebra agrees closely
with the Porto Rican series.
The discovery of this form in Porto Rico proper greatly reduces the
differentiation of the Virgin Island fauna.

Eleutherodactylus brittoui* Seihmidt
Text Fig. 18S
Eleu1liecriidairly/lu brifltoi NSehhindt, 1920), Ann. N. Y. Acad. Sci. Vol. XXVITI,
p. 179, Fig. 7.
No native name is available for this species.
Type locality.-El Yunque, near the Forester's Cabin, about 1300 feet
altitude, Luquillo Forest Reserve, Porto Rico.
Distribution.-Known only from El Yunque and Maricao.
Specimens rollected.--I : El Yunque and Maricao.
Diagnosis.-Derivedl from Eleutherodalitylus antilleusis, from which it
is distinguished by its small size, its sharp eanthus rostralis, which is
continued as a dorso-lateral angle some distance behind the ear, and its
more posteriorly placed nostril.
Original descriptioi.--"Hlabitus slender, head narrower than the body.
legs rather short, snout sharp-pointed; nostril two-fifths the distance
from the end of the snout to the orbit: canthius rostralis sharp: inter-
orbital space broader than the eyelid : heel reaching the anterior border
of the orbit : heels meeting but not overlapping when the legs are at right
angles to the body; top of snout flat, as is the anterior half of the back
behind the eyes, the side of the body being vertical anteriorly; vomerine
teeth in two small rounded patches, behind and within the choanna:
tympanmn indistinct, separated from the eye by less than its diameter;
dorsum smooth, venter coarsely granulate; digital disks small, as long as
wide: a well-defined tarsal fold : a well-developed subgular vocal sac.
"Dorsumn light grayish brown, venter lighter. Two black spots between
the eves, one on the middle of the back. and three posteriorly on the
back, above the groin: legs with a single faint darker bar on the femur:
concealed surfaces or the femur not reticulated: a black subeanthal
streak, continued below the dorso-lateral angle behind the eye."

Named for Dr. Nathaniel IL. IBritton. Chairman of the Committee for the Rcientific
Survey of Porto Rico and the Virgin Islands. New York Academy of Sciences.








SCIENTIFIC SURVEY OF PORTO RICO


MEASURE-M ENTS
Tip of snout to vent .............................................. 16 n nm .
Tip of snout to posterior edge of tympanum ........................ (i
Greatest breadth of head........................................ 6
Foreleg from axilla ............................. ............... 9
Hind leg from vent............................ ................ 23
Tibia ............................... ............ .............. 8

Remarks.-The three paratypes are closely similar in size and struc-
tural characters to the type. Two have the black subcanthal and shoulder
mark outlined with white above. One lacks the dorsal black spots.
The single specimen from Maricao was taken singing in herbage along
the roadside, together with E. portoricensis and E. antillensis. Two







I Fi. 18.-Elealthcrodactylus brittoni. A. 31.
N, II. No. 10318. Twice natural size.








were taken singing on El Yunque, likewise in low herbage, and the last
was found by accident in collecting E. wight ianae.
The song of this species is a succession of clicks, less shrill and less
rapid than in E. gryllus.
This species stands in the same relation to E. antillensis a, E. grE;it. s
does to E. porloricensis.

Eleutherodactylus wightmanae* Schmidt
Text Fig. 19

Eleuthicrodachtyls wiglhtinanac Schmidt, 1920, Ann. N. Y. Acad. Sci., Vol.
XXVIII, p. 181, Fig. S.

No native name exists for this species.
Type locality.-El Yunque, near the Forester's Cabin, about 1300 feet
altitude, Luquillo Forest Reserve, Porto Rico.

Named for the anutor's wife, Margaret Wightman Schmidt, whose loyal assistance
contributed largely to the success of the work in Porto Rico in 1919.








S'CMIDT, AMPHIBlANS OF PORTO RICO


DisIribution.-Known from El Yunque and Maricao, at opposite
ends of the island.
Specimens collected.-13: El Yunque and Maricao.
Diagnosis.-Size small; snout pointed; nostril much nearer to the
tip of the snout than to the eve; tympanum small, distinct, separated
from the eye by about its own diameter; vomerine teeth in two straight
series, in the same line, extending as far laterally as the choante, and
about the diameter of a choana behind them ; toes free, digital disks well-
developed; tibio-tarsal articulation reaching the anterior border of the
eye; heels overlapping when the legs are placed at right angles to the
body; skin rugose above, with elongate folds and ridges; venter rugose;
thighs granular.
Original description.-"Head as long as broad, narrower than the
body: snout pointed, its length anterior to the eyes once and a half the







FIG. 109.-Eleutherodactylus wiight- "
manae. A. 'M. N. H. No. 10220. '1
Twice natural size. t







interorbital width: nostrils one-third the distance between eye and tip
of snout from the latter: tympanum distinct, small, about one-third
the diameter of the eye, separated from the eve by a little more than its
own diameter; canthus rostralis sharp; elbow and knee pressed along
sides overlap: heels overlap when the legs are placed at right angles to
the body: tibio-tarsal articulation reaching the anterior border of the
eye; disks of fingers and toes well-developed: digits slender, free; first
toe distinctly shorter than the second; no tarsal folds; vomerine teeth in
two straight series, separated in the median line, extending laterally as
far as the outer border of the choanae, and about the diameter of a
choana behind them; tongue large, slightly nicked behind; skin rugose
above, with longitudinal lines or folds, the most distinct of which orig-
inate behind the orbits and extend backward about two-thirds the length








,sIENTIFIC SURVEY OF POUTO RICO


of the hack: a less distinct mid-dorsal ridge from snout to vent: venter
and outer face of the thighs moderately rugose: a subgular vocal sac.
"Brown above, with a black suheanthal line. extending over the ear
half way along the sides; a black spot on each side of the back over the
groin; venter uniformly light; a single dark crosshand on the radius ; one
on the femur, tibia, and tarsus (in line when the legs are folded), and a
dark spot on the metatarsns; anterior and posterior faces of the thigh
dusky."
1 AS'A SUREMEN TS
Tip of snout to vent............................. .............. 20 mrm.
Tip of snout to posterior edge of tympanum ...................... 7.5
Greatest breadth of head ......................... ................ 7.
Foreleg from axilla ............................................. 11
Hind leg from vent........................... ................ 30
Tibia .............................. ............. .............. 10

Remark's.-In structural characters the twelve paratypes agree closely
with the type. Two specimens are light gray, instead of brown, with
only indications of the black spots; in most specimens the postocular
dark streak is broken up into a series of spots: one specimen is light
brownish gray on each side. the area between sharply darker: the bars on
the legs are distinct in all specimens.
The plaintive, diminuendo note of this small species is one of the most
characteristic sounds in the amphibian chorus of the Luquillo Forest.
Its song consists of a series of six or eight whistled notes, each slightly
lower in pitch and a little fainter than the previous one. The creature
sings habitually on the ground or in the lowest leaves of plants. To lo-
cate its position from its song it is particularly difficult, partly because
it is usually well concealed, partly on account of the peculiar ventriloquy
of its voice.
Nothing has been added to our knowledge of this species since its dis-
covery.
Eleutherodactylus richmondi Stejneger
Text Figs. 10 and 20

Eleitherodelctylis riiinonidi Stejneger. 1904, Rept. U. S. Nation. Mus., 1902.
p. 593. Figs. 25-29.-Barbour. 1914. 2Mem,. Mus. Comp. Zool., Vol. XLIV,
p. 247.-Schmidt, 1920, Ann. N. Y. Acad. Sci., Vol. XXVIII, p. 183. Fig. 9.

No native name exists for this species.
Type locality.-Catalina Plantation, about 890 feet altitude, eastern
slope of El Yunque, Porto Rico.








SCHMIDT), AMPHIBIANS OF PORTO RICO


Distrib ution.-Known only from Porto Rico and apparently ciiifined
to El Yunque.
Specimens collected.-11 : El Yunque, from 1300 feet to the peak.
Diagnosis.-"Toes free without a vestige of web; belly smooth; tym-
panum distinct, less than one-half the diameter of the eye; vomerine
teeth in two long angular transverse series, extending beyond the
external border of the inner nares and some distance behind them;
digital disks small; nostril much nearer tip of snout than eye; hind limbs
not cross-barred."
























FIG. 20.-Eleutherodactylus richmondf. A. M. N. H. No. 10237. Twice natural size.

Original description.-"Tongue narrow, somewhat emarginate behind;
vomerine teeth in two angular series behind the choanm, their distance
from the ehoanme greater than the diameter of the latter; inner arm of
each vomerine series longer, outer extending laterally beyond the ehoanem;
nostril much nearer the tip of the snout than the eye, the distance
from the eye slightly less than the diameter of the latter; upper eyelids
somewhat narrower than interorbital space, tympanum slightly less
than one-half the diameter of the eye, its distance from the latter
slightly less than its diameter; disks of fingers rather small, first finger
shorter than second; disks of toes small, first toe short, only reaching








SCIENTIFIC SURVEY OF PORTO RICO


subarticular tubercle of second; subarticular tubercles well devel-
oped; two well developed metatarsal tubercles; no plantar tubercles;
no tarsal fold; the bent limbs being pressed along the sides, knee and
elbow overlap; hind limb being extended along the side, heel reaches
center of eye; hind limbs being placed vertically to the axis of the
body, the heels overlap; skin above and on flanks granular, underside
smooth; posterior aspect of femur areolate."
Remarks.-The coloration in life has been described by Stejneger as
follows:-"Back dusky chestnut, lighter on sacrum; from each nostril
along canthus rostralis, edge of eyebrow and sides of back a narrow dirty
bluish-white stripe somewhat wider on sides of back than on canthus
rostralis; sides of face and flanks below this stripe blackish, legs black-
ish: fore legs marbled with pale drab, hind legs with dull pale chest-
nut; under side dull greenish gray, with an ill-defined yellow spot in each
groin, and marbled with dusky brown on throat and under side of hind
legs. Iris blackish, brassy above pupil."
Like the larger series examined by Stejneger, the specimens collected
on El Yunque by myself are extremely uniform in structural characters
and in coloration. The only variation noted is the occasional lightening
of the chestnut color of the dorsal area between the light dorso-lateral
lines. The proportions are quite different in this species from the other
Porto Riean species of the genus:-
A.M.N.H. U.S.N.M.
Parts mnpasured No. 19233 No. 26884
Tip of snout to vent.................................. 32 mm. 3S mm.
Tip of snout to posterior edge of tympanum............... 1:1. 15
Greatest breadth of head................ ........... 12 15
Foreleg from axilla ...................................... 21 24
Hind leg from vent ................................ 51 62

Two extremely small specimens, measuring 9 and 11 mm., respectively,
probably are recently transformed. They are colored like the adults.
All of the specimens known were found under stones or. palm leaves
on the trail or on damp ground, associated with E. porloricensis. The
males of this species were not discovered singing and its voice is un-
known. The slender digits and small adhesive disks probably indicate
that it is more terrestrial in its habits, and the eggs may prove to be laid
on the ground. like those of E. luteolus of Jamaica.
This species is allied by the form of its vomerine teeth to E. lentus of
the Yirgin Islands, E. monensis of Mona Island, E. weinlandi of His-
paniola and perhaps to E. jamaicensis of Jamaica. This group of species
thus composes an interesting series of vicariating forms.








SCHMIDT, AMPHIBIANS OF PORTO RICO


Eleiitherodlatylus inonensis (Meerwarth)
Text Fig. 21
HIIylodes moncnsis Meerwarth, 1!01, Mitt. Naturh. Mus. Hamburgh, Vol.
XVIII, p. 39, I'l. 1 (Fig. 11), P1. 2 (Figs. 4-5).
Eleutherodactyluht monensis Stejneger, 1904, Rept. U. S. Nation. Mus., 1902,
p. 595, Figs. 30-34.-Barbour, 1914, Mem. Mus. Comp. Zool., Vol. XL1V, p.
247.-Schmidt, 1926, Publ. Field Mus. Nat. Hist., Zool., Vol. XII, p. 154.

No native name exists for this species other than "ecoqui.'"
Type locality.-Mona Island, West Indies.
Distribution.-Confined to Mona Island.
Specimens collected.-41: Mona Island.
Diagnosis.-Yomerine teeth in long arched transverse series behind the
choanme; belly smooth; soles of feet tubercular; hind foot nearly as long











FIG. 21.-Elitherodactlis mo-
,ensis. A. M. N. H. No. 24463. /












as fore leg; color of back and sides pale, with brownish markings.
Original descriplion.-"This species is closely allied to the previous
one [Hylodes lentus Cope] and differs from it in the following features:
in II. lentus the femur, measured from the ischio-pubic crest to the edge
of the knee is shorter than the tibia and at most as long as the distance
from axilla to groin, while in H. monensis it is as long as the tibia and
longer than the distance from axilla to groin; the tongue is wedge-
shaped; the vomerine teeth are also in two rows behind the choanm, but








SCIENTIFIC SURVEY OF PORTO RICO


the angle in each row is more obtuse, and the outer portion (beyond the
ankle) is about half as long as the inner, instead of subequal, as in II.
lent us.
"The color of the belly is uniform whitish yellow, that of the upper
side a gravish flesh-color with small brown spots sparsely distributed on
the back, sides, and limbs, and a more or less well-defined star-shaped
figure formed from larger brown spots between the shoulders. A brown
line extends from the nostril to the eye."
Remarks.-This species is well characterized in the original descrip-
tion and figures. Thlie series of specimens secured by Anthony in 1926
agrees with the Hamburg originals and the single specimen in the
National Museum described in detail by Stejneger.

Eleutherodactylus unicolor Stejueger
Text Figs. 22 and 23

Elentlherodactyluus isnicolor Stejueger, 1l9i04, Rept. U. S. Nation. Mus.. 1902, p.
.0 7. Figs. 35-39.

No native name.
Type localily.--'amp on El Ynnque at 2978 feet altitude, Luquillo
Forest Reserve, Porto Rico.
Distribution.-Known only from the type locality.
Diagtnosis.-"Toes free without a vestige of web; belly granular;
tympanunm distinct, one-third the diameter of eye: vomerine teeth in

F FIG. 22.-Eleathecrodactylus uni-
C'A /color. Side of head (left), top
V>-- of head (center), and Inside
of mouth (right) of type.
(After Stejneger.)


two short, straight series, not extending beyond the inner nares; head
not broader than body; interorbital space equals upper eyelid: upper
surface smooth; second finger longer than first; inner metatarsal
tubercle large; digital disks small; nostrils intermediate between eye
and tip of snout: hind limbs not cross barred."
Original description..-"Tongue medium, oval, entire behind : vomerine
teeth in two short straight series behind the choanm, but not extending
laterally beyond them, widely separated in the middle; snout declining
rapidly from the eyes to the tip; nostril situated about halfway between
eyes and tip of snout, their distance from the eyes one-half the diameter
of the eye; upper eyelids as wide as interorbital space; tympanum small,


. 6








Sf'HMIDT, AlMI'llIIIANS OF PORTO ICO


about one-third the diameter of the eve and distance from the latter
more than its own diameter: fingers with exceedingly small disks, first
slio'htly shorter than second : disks of toes better developed, first toe much
shorter than second subarticnlar tubercles well developed ; no plantar
tubercles: two well-developed metatarsal tlubereleo : no tarsal fold; hind
limbs being' bent forward, heels reach the ears, bent vertically to the
axis of the body., the heels do not tonch: skin above, throat, chest, and
anterior aspect of femurs smooth: belly and sides granular."


FIG. 23.--Eleatherodactylus unicolor. U. S. N. M. No. 260063, type.
(Courtesy of Dr. Leonhard Stejneger.)


Twice natural size.


Remarks.-Stejneger describes the coloration in life as follows: "Uni-
formly dusky chestnut above and below, with scattered, scarcely visible
pale dots; a short postocular dusky band descending behind the tym-
panum."
3IEASUREMENTS OF THE TYPE
Tip of snout to vent........................... ................ 16.5 m m .
W idth of hend .................................. ........... .... 6
D iam eter of eye.......................... .................... 2.5
Diameter of tympanum ........................................ 0.8
Length of arm ................. ............................. 7
Length of leg......................... ........ ............... 22


F








SCIENTIFIC SURVEY OF P'ORTO RICO


This species is still known only from the type. Its characters are so
peculiar that additional specimens are much to be desired. Its pectoral
girdle should be examined.

Cl ss REPTILIA

Order SQUAMATA

The Squamata comprise the lizards and snakes, corresponding to the
two suborders Sauria and Serpentes. The necessity of associating these
two groups as a single order is reflected by the difficulty in framing a
non-technical distinction between them. Anatomically they may best
be distinguished by the separation of the mandibles anteriorly in the
snakes, while in the lizards the mandibles are united by a suture. The
limbless lizards of the family Amphisbaenidae and the blind snakes of
the family Typhlopidae offer the chief difficulty in distinguishing the
lizards and snakes in Porto Rico. The Amphisbaenidae, however, are
immediately distinguishable by the absence of ordinary overlapping
scales, the skin being divided into rectangular segments in regular rows
and rings.
Suborder SAURIA

KEY TO THE GENERA OF PORTO RICAN LIZARDS

A. Four limbs present.
B. Top of head covered with numerous small scales or granules.
C. Eyelids vestigial, not closing; pupil vertical........... (Gekkonidae)
D. Toes dilated at the base, the terminal joints free, claw-shaped
(see Fig. 24, left) ..............................Hemidaetylus
DD. Toes dilated only at the tip, which is provided with a circular
plate beneath (see Fig. 24, right)..............Spihaerodactylus



FIG. 24.-Digits of Hemidaetylus maboula (left)
and Sphaerodactytls mnacrolqpis (right) con-
S( trasted; (From Stejneger.)





CC. Eyelids well-developed, functional...................... Iguauidae
D. Toes dilated at base, tip claw-shaped, as in Hemidactylus; no
"combs" on toes: an extensible throat fan in the male, indicated
in the fem ale........................................... Anzotis
DD. Toes not dilated, "combs" present; no throat fan..... Cyclura









SCHMIDT, AMPHIIIBIANS OF PORTO RICO


BB. Head with large regular shields above.
C. Occipital shield present; limbs relatively small (Anguidae) .. Cclestus
CO. Occipital shield absent, limbs moderate or well-developed.
D. Ventral scales not imbricate, in regular longitudinal and trans-
verse rows (Teiidae) ..................................Ameira
DD. Ventral scales imbricate, in oblique rows (Scincidae) Mabuya
AA. No limbs; eyes merely indicated by a dark spot beneath the skin, which
is divided into small rectangular areas on the body (Amphisbaenidae)
Amphisbaen a

GEKKONIDAE

Hemidattylus Oken

Hemnidactylus nmabouia (Moreau de Jonnus)
Text Figs. 24 and 25

Gecko mabouia Moreau de Jonnes, 1818, Bull. Soc. Philom., Paris, 1818, p.
138; 1821, Monogr. du Geeko mabouia, p. 1.
Hemidactylus imabouia Dumeril & Bibron, 1836, Erp6t. GWn., Vol. III, p.
362.-Dumeril, 1851, Cat. Method. liept. Mus. Paris, Vol. I. p. 39.-Rein-
hardt and Luetken, 1863, Vid. Meddel. Naturh. Foren., Copenhagen, 1862,
p. 174, p. 275.-Cope, 1868, Proc. Acad. Nat. Sci., Phila., p. 311.-Boulenger,
1885, Cat. Lizards Brit. Mus., Vol. I, p. 122.-Strauch, 1887, Mem. Acad.
Sci. St. Petersb., (7), Vol. XXXV, No. 2, p. 31.-Carman, 1887, Bull. Es-
sex Inst., Vol. XIX, p. 18.--Meerwarth, 1901, Mitt. Naturh. Mus. Ham-
burg, Vol. XVIII, p. 17.-Stejneger, 1904, Ann. Rept. U. S. Nation. Mlus.,
1902, p. 599, Figs. 40-45.-Wolcott, 1924, Journ. Rept. Agric. Pto. Rico, Vol.
VII, p. 13.
Hemidactylus mabuya Fitzinger, 1843, Syst. Itept., p. 105.
Hemidactylus mabuia Cocteau, 1843, in Sagra, Hist. Fis. Pol. Nat. Cuba, Hist.
Nat., Vol. IV, p. 95, Pl. 16.-Gundlach, 1867, Rept. Fisico-Nat. Cuba,
Vol. II, No. 5, p. 12; 1875; An. Soc. Espaii. Hist. Nat., Vol. IV, p. 358;
1881, idem. Vol. X, p. 308.-Boettger, 1893, Knat. Rept. Mus. Senckenlnerg,
Vol. I, p. 28.

Type locality.-St. Vincent, West Indies.
Distribution.-The West Indies, except perhaps the northern Lesser
Antilles; northern South America. As I have remarked above, I do
not consider the African geckos usually referred to this species as con-
specific with the West Indian form.
Diagnosis.-A gecko of moderate size, the basal half of each digit ex-
panded and provided beneath with pairs of lamellae, the distal half com-
pressed, arising from within the tip of the expanded portion, curved,
and provided with a claw at the tip; enlarged dorsal tubercles rather
widely spaced, convex, not distinctly keeled; a long series of femoral
pores, scarcely interrupted on the mid-line.








,N'I-A\ TIFIf' SURVEY OF PORTO RICO


Original descripiotn.-"Digits expanded throughout their length, with
two rows of transverse lamellae beneath; each terminating in a hooked
claw: the back studded with tubercles and the tail with spinose scales;
transverse plates oii the under side of the tail: femora with pores be-
neath."
Remarks.-This species is the only one of the larger geckos found in
Porto Rico. It is curiously scarce in collections and, if as rare in fact as
this scarcity would seem to indicate, it may not he an indigenous mem-
ber of the Porto Rican fauna.
Nothing is known of the habits of this species in Porto Rico beyond
the fact that it has been taken in San Juan and that it was reported by






VIi;. 25i.-Head of Hemidactlctilu s mnbouii. (From Stej negi'r.l







Dr. E. Giraywood Sinvtl of the Agricultural Experiment Station at Rio
Piedras to frequent the vicinity of the are-lights in Rio Piedras at night
to prey upon the insects.
Beyond this meagre information nothing is known of its distribution
on the island. In the West Indies as a whole it is widespread-from
Trinidad and Barbadoes to Cuba and .lamaica. It is curious that it has
not been recorded from the northern group of smaller islands in the
Lesser Antilles. I have little doubt of its identity as a species through-
out this range. A West African species, Hiemidactylas brookii, occurs
in Hispaniola (Port-au-Prince), where it was doubtless introduced by
the slave-ships. The Mediterranean Hemidartylus turcicus, it is said.
has become established at Key West.

Sphaerodactylus Wagler
Spha erodactylus miarrolepis G(inuther
Text Figs. 24 and 26
Salaniaidra. Salanimndrita: Salainaiquna. Salamniquita; Lucia (?) ; Ranit ;
Lagortija eaheza d(e muerte








SCI11IDT, AMPHIBIANS OF PORTO RICO


Sphacroduictyls macro(lpiS Giinther, 1859, Ann. Mag. Nat. Hist.. (3). Vol. IV.
Ip. 215, Pl. 4, Fig. B.- Blarbour. 1 .14, Mem. AMus. Comp. Zool., Vol. XLIV,
p. 270.: 1115. P'rnc. Biol. Soc. Wash., Vol. XXVII. p. 72: 1917, Vol. XXX.
p. 9S.- Schmidt. 1920, Ann. N. Y. Acad. Sei., Vol. XXVIII. p. 1S4.-Bar-
lihour. 1921, Mem. 3ils. Co omp. Zool., Vol. XLVII, p. 253. I'l. 6, Figs. 2-3,
I'. 19, Figs. 5-S.-VWolcott, 1924, Journ. Dept. Agrie. 1'to. Rico, Vol. VII,
I). 1:t.
Splhtrodacl/s,' n maroloh',pi, moiniin i.sN Meerwarth, 190)1, Mitt. Naturh. Mus.
Hamlburg. Vol. XVIlI. p. 20.
Sphrtriwh ciiiti n/ll. oiiti 'ix Stejueger, !04, Rept. U. S. Nat. MIns., 1P02, p.
G07.-- arbour, 1 914, Meim. Mus. Comp. Zool., Vol. XLIV, p. 270.
Sph(rc-od(1ctylus grundinsquiomis Stejneger, 1904, Rept. U. S. Nat. Mus.. 1902,
p. 602. Figs. 4l6-52.--Barliour, 1914. Alem. Mus. Comp. Zool., Vol. XLIV,
p. 270.-Fowler, 1918, Publ. Carnegie Inst. Wash.. No. 252, p. 7.

The species of this genus are usually known as Salamandritas (or
Salamanquitas) to the Spanish-speaking West Indians. The wriggling
mode of progression of these tiny geckos is quite salamander-like and
the name is to that extent appropriate. Stejneger found that the name
"Lucia" or "Santa Lucia" was applied to this species hby the children in
Luqnuillo. This, however, must have been an unfortunate localism, as
these names are used everywhere in Porto Rico for the Moabuya, a totally
different creature. The equally local use of the name "lanita" at Aibo-
nito illustrates the tendency to apply a name which exists in the language
to any unidentified species.
Type locality.-St. Thomas. Virgin Islands.
Distribtiomn.-Widespread on Porto RIico, where it is recorded from
Aibonito, Bayamon, Cantailo, Coamo Springs. Ensenada, Luquillo,
Maricao, Ponce and El Yunque. It is evidently not confined to the
coastal plain as supposed by Stejneger. On the outlying islands it is
known from Mona and Vieques and occurs on most of the Virgin Islands.
Specimens collected.-45, from Aibonito, Bayamon, Catafio, Coamo
Springs, Ensenada, Maricao, El Yunque and Mona Island.
Diagnosis.-A small geckoid lizard with unexpanded digits which are
provided with an enlarged flat circular scale beneath the tip: dorsal
scales keeled, imbricate, somewhat variable in size; no vertebral series of
small scales.
Original dlescriplioa.-"Body surrounded by about forty longitudinal
series of scales of rather large size; no vertebral streak of smaller ones,
those of the back keeled, of the belly smooth. Trunk and tail uniform
blackish brown, in younger individuals some scales with blackish tips:
head greyish brown, marbled with black; jaws and throat striolated with
blackish.








SCIENTIFIC SUkVEY OF PORTO RICO


"The snout is of moderate extent, and slightly pointed; all the upper
surface of the head and the sides are covered with scales of moderate
size; there is an exceedingly small horn-like spine above the middle of
the orbit. The rostral shield is low, and bent backwards on the upper
surface of the snout; the sides of the jaw are margined with three elon-
gate labials; the nostril is situated above the posterior extremity of the
rostral shield and first labial, and exceedingly small. The anterior
lower labial is single; a series of three other shields covers the lateral
margin of the lower jaw. The scales of the throat are small, those of the
breast and of the extremities keeled. The ear-opening is very small,
one-third only of the width of the eye. The fingers and toes have an en-






Fir. 2G.-Head and shoulders of
S Sphaerodaetylus macrolepis. A.
M. N. H. No. 13037 (A) and
S No. 13697 (B). showing two
S common types of pattern.
Two and a half times natural

if





tire and unarmed disk. The tail is covered with smooth scales, rather
smaller than those of the trunk; there is a series of larger ones, plate-
like, along the lower medial line. No femoral or anal pores.
"I add to the statement of the coloration given above that the belly is
uniform dirty white, and the tail minutely dotted with blackish. Two
specimens were in the collection."
Remarks.-Barbour (1917, p. 98), after examining a considerable
series of Spaerodaclylus macrolepis from the Virgin Islands, expressed a
measure of doubt as to the distinctness of S. grandisquamis. Both Bar-
bour and I have subsequently reached the conclusion that it is untenable.
Stejneger separated S. grandisquamis and S. monensis from S. macrolepis
solely on the size of the scales, which he gives as 34-38 about the body in
S. grandisquamis, 4G6-48 in S. monensis. In the series from Porto Rico
collected by myself the variation is as follows:








SCHMIDT, AMPHIBIANS OF PORTO RICO


Scales about the body 32 36 40 44 48 52 56
Number of specimens 2 8 3 4 5 G 2

In five specimens from Mona Island the number of scales varies from
44 to 52; as S. macrolepis is intermediate between S. grandisquamis and
S. monensis, it is evident that the variation in the present series includes
all three supposed forms. There is probably a somewhat different range
of variation on the several islands but the extremes are certainly in-
cluded in that of the Porto Rican series.
Reproduced tails have a much widened series of median ventral scales.

MEASUREMENT OF A. M. N. H. No. 13324
Total length ................................................... . 66 m m .
T ail ............................................................. 34
Tip of snout to posterior edge of ear............................. S
Breadth of head........................ ........................ 5.5
Foreleg from axilla............................ .... ............. 7.5
H ind leg from groin...................... ..................... 10

The smallest specimen measures 12 mm. from snout to vent.
This species is highly variable in coloration. The absence of the elab-
orate head pattern appears to be due to the general darkening of the
coloration, as all of the lighter specimens have it in some form or other.
Only two specimens are without a trace af the black shoulder band with
its two white spots. In the remaining specimens its development is very
irregular, the white spots persisting even when the black band is indis-
tinguishable. The darker dorsal spots are usually in rows, and in some
specimens form longitudinal lines. The throat is uniform or heavily
dotted with dark spots. Juvenile specimens are usually dark in color,
with the white spots of the scapular band distinct. One specimen (No.
13811) is light brown in ground color with five light gray longitudinal
stripes; one, median from between the eyes to the base of the tail; two,
dorso-lateral from the upper posterior corner of the eye to the base of the
tail, and two, lateral from the eye through the ear to the groin, the latter
only narrowly separated from the light venter. In specimens from
Mlona Island, the scapular black band is larger, outlined with light
color, the brilliant white spots transverse, forming a continuous line in
one specimen.
Habits.-The usual habitat of this species is the ground cover of dead
leaves in coffee plantations and forest. Elsewhere it is found under stones
and logs. Along the base of the limestone cliffs on the hills back of
Catafio, Sphaerodactylus were numerous, scurrying for the crevices at the
base of the cliff when alarmed. One specimen was found under a leaf








SCIEAT'FI(C SURVEY OF PORTO RICO


sheath, and others on the trunks, of banana plants. The specimens from
Bayamon were found in picking up rubbish on freshly cleared land.
These reptiles frequently venture forth in broad daylight, but are prob-
ably essentially crepuscular or nocturnal.
Four of the specimens collected on Mona Island were taken among the
limestone boulders on tile west side of the island, and of these three were
secured at night with the hand lamp. Two were taken beneath pieces of
coral on the flat terrace on the south side.
Their extreme agility makes them difficult to catch, and the tail is
often broken or the skin torn. When caught, they frequently turn a
uniform light gray, becoming brown again in the collecting bag.
An egg, probably of this species, was found under a log at Aibonito,
August 21, 1919. It is white, discolored by stains, with a hard and
smooth shell, 6 x 4.5 mnm.

IGUANIDAE

Anolis Daudin

The lizards of the genus Anolis, characterized by expanded digits that
have a raised terminal claw-like portion, and by the presence of a throat-
fan, usually brightly colored, which may be distended vertically, are the
most ubiquitous of West Indian lizards. The multitude of species in
the West Indies is paralleled in Central America and in northwestern
South America. Wherever lizards of the genus Anolis occur, a few
species are usually extremely abundant-thus Anolis crislatellus on Porto
Rico. A. cybotes on Hispaniola, A. sagrei on Cuba and at Belize.
The power of color change is highly developed in the Anoles, and they
are frequently miscalled "ehamaeleons" or "cameleones." As Stejneger
remarks, "Anolis" might well be adopted as a vernacular name. In spite
of their color-change, the species are almost invariably distinguishable
byv some feature of their coloration, and I have drawn up a key to the
males of the species on this basis to supplement the synopsis based on
structural characters devised by Stejneger, which must be referred to
when the color characters fail.
KEY TO TIlhE SECIEs OF ANOLIS RECORDED FROM PORTO RICO
A. Dorsal scales entirely separated from each other by several circles of
granules; size large: male with tail crest......................A. euvieri
AA. Dorsal scales juxtaposed or imbricated: size moderate or small.
IS. Dorsal scales (all. or with the exception of two rows on the median
line) granular or tubercular. differing but little, if at all, from laterals,
but very much from the much larger ventrals. which are smooth or
feebly keeled.










Sf'HMIDT, AUP'JHI/IAN OF PORTO RICO


C. Two. or more, shields or scales between the superciliaries and the
supra ocular semicircle bordering the sIIp rai nl-hir granules anteriorly.
Tail of male crested.
10. Snpraocniar semicircles separated Iby at least two scale rows:
occipital shield separated from suprin(iular semicircles by at
least five scale rows (Fig. ) ...................... 1. gyundluchi
I)r. Supraocular seminircn-les in cointalct or with at mllost a single
series of scales between: occipital shield separated from
snll raocula rI selmicireles by at most four scale rows ..........
-1. cris/atlfellux
CC. One shield between the superciliaries and the supraocular semi
circle bordering the siupraocnlar granules anteriorly.
I). Width of head as great. or greater than distance froin tip of
snout to center of eye: anterior femoral scales keeled, gradually
diminishing : color greenish ....................... A.. rerimanni
DD. Width of head less than distance from tip of snout to center
of eye; anterior femoral scales smooth, abruptly larger than
the others; color brownish or grayish............... .slraulus
BB. Dorsal scales large, flat, keeled. imbricate. very much like the ventrals.
which are very strongly keeled. the keels forming continuous ridlres.
C. Lateral scales granular.
D. Width of head muchli more than half the distance from tip of
snout to ear-opening; four to six inedian dorsal scale rows more
or less abruptly larger than the others; skin of dewlap in male,
orange ......................... .. ................... rugi
DDl. Width of head about mone-half the distance from tip of snout
to ear-opening; dorsal scales gradually increasing in size from
the laterals toward the median rows; skin or dewlap in male.
crim son ....................... ........... .. I pulchellus
CC. Lateral scales imbricated, keeled ..................... ponicensis

COLOR KEY TO P'ORTO RICAN ANOLES

A. No longitudinal stripes.
1. Color brown or gray, never bright green.
C. No short transverse saddle-shaped vertebral spots.
1). Iris dark brown ; skin of dewlap greenish yellow, its edge brown-
ish orange...................................... A. c'rix tellu.s
DD. Iris metallic blue: skin of dewlap orange olive, with distant
yellow scales.................................. gundlaclhi
CC. Short black saddle-shaped spots on the mid-line of the back: dew-
lap bright orange. .................................. straltulus
BB. Color bright green (if not green. no transverse bands or spots).
C. Size large. front of head flat, bony........................ A ncrier
CC. Size moderate, front of head concave................A cccriaeni1
AA. Light longitudinal stripes present.
B. Throat-fan white..................................... A. ponccnsis
'B Throat-fan crimson................... ............. 1. ulchcllaus
BBU. Throat-fan orange..................... ............... A. krugi









SCIENTIFIC SURVEY OF PORTO RICO


0



~,iJ







/


FIG. 27.-Heads of Porto Rican Anolis. Anolls cu ier! (left of top row), Anolis crista-
tellus (center of top row). Anolis gundilochi (right of top row) ; Anolis evermanni
(left of middle row), Atolis stratulus (center of middle row), Anolis krugi (right of
middle row) ; Anolis pulchellus (left of bottom row), Anolis poncensis (right of bot-
tom row). (From Stejneger.)








SCHMIDT, AMPHIBIANS OF PJORTO RICO


Anolis cuvieri Merrem
LI'garto, Chipojo
Text Figs. 27 and 28
Jnolis curicri Merrem, 1820, Syst. Amphib., p. 15.-1ioulengir, 1S85, Cat.
Lizards, Brit. MIns., Vol. II. p. 23.-Garman, 18S87, Iull. Essex nlst.. Vol.
XIX, p. 27.-Stejueger, 1904, Rept. U. S. Nation. Mus. 19(12. p. 627, Figs.
81-S4, 87.-Barbour. 1914. Monm. Mus. Colip. Zool., Vol. XLIV. p. 273.-
Fowler, 191S, Papers Dept. Marine Biol., Carnegie Inst., Vol. XII, p. 7.-
Shmnidt, 1920, Ann. N. Y. Acad. Sci., Vol. XXVIII, p. 1S3.-Smyth, 1920,
Rev. Agric. Pto. Rico, Vol. IV, p. 19.-Wolcott, 1924, Journ. Dept. Agric.
Pto. Rico, Vol. VII, p. 14.
Anolius velifer Cuvier, 1829, RWgne Anim., 2nd Ed., Vol. II, p. 29, Pl. 5,
Fig. 1.-Guerin, 1830, Icon. Regne Animi., Rept., Pl. 12, Fig. 1.
Anolis velifer Dumnril and Bibron, 1837, Erp6t. G6n., Vol. IV, p. 164.-Dumeril,
1851, Cat. Method. Rept. Mus. Paris, Vol. I, p. 59.-Reinhardt and
Luetken, 1863, Vid. \Ieddel. Naturh. Foren., Copenhagen, 1862, p. 260.-*
Cope, 1868, Proc. Acad. Nat. Sci. Phila., p. 312.-Peters, 1876, Monatsber.
Akad. Wiss. Berlin, p. 705.-Gundlach, 1881, Anales Soc. Espafi. Hist.
Nat., Vol. X, p. 308.-Stahl, 1882, Fauna Puerto-Rico, p. 69, p. 159.
Xiphosurus relifcr Cope, 1861, Proc. Acad. Nat. Sci. Philn., p. 208.

Type locality.-Jamaica (erroneously).
Distribution.-Anolis cuvieri has been taken at Aibonito, Catalina
Plantation (El Yunque), Ciales, Humacao, Luquillo, Mayagiiez and
Utuado. It is probably not found in the arid southwestern corner of
the island, but ranges quite generally over the remaining part of Porto
Rico. It is recorded from Vieques and Tortola of the Virgin Islands.
Its absence from the other Virgin Islands is perhaps due to difference
in the habitat conditions or perhaps to extinction. It is nearly allied
to Anolis ricordii of Hispaniola.
Specimens collected.-11. Aibonito.
Original descriptioni.-"The rayed fin extending from the base to the
middle of the tail, with twelve to fifteen rays. Throat fan extending to
the breast."
Remarks.-Stejneger's discussion of the historical aspect of the
taxonomy of this species, and of its relations with the IIispaniolan Anolis
ricordii is a model of completeness and clarity. His description of a
male specimen follows:
"Top of head flat, with only shallow depressions on prefrontal and
occipital region, the scales being rather small and roughly keeled and
tuberculated, even those on top of the snout, but especially those of the
supraorbital semicircle and frontal ridges; about nine enlarged supra-
oculars. flat, keeled, and in contact with the semicirculars: supraorbital








sIEXT'IF1C SURVEY OF POQITO RICO


semicircles separated by about three scale rows from each other and from
the occipital, which is barely noticeable: scales surrounding the occipital
depression on the sides and behind rather large, flat, polygonous. each
with a strong keel: six loreal rows, the scales composing the lower row
next to the supralabials largest; one row of large keeled suboculars;
7-8 supralabials to under the center of the eye: temporals flat, with a
low tubercle, all the scales of the sides of the head being more or less
rugose or wrinkled: ear-opening rather small, upright, oval; back and



















FIG. 28.-Caudal crest of Anolis Cnrieri (left), of A. yundlachi (center), and
A. crislatellus (right).

sides covered with uniform scales tuberculated or keeled, separated from
each other by one or more rings of minute granules; on the median line
of the neck and back a series of about fifty triangular spines forming a
saw-tooth ridge scarcely connected with the caudal crest; ventral scales
about same size as dorsals, though more closely set, but not -keeled or
distinctly tuberculated except on the flanks; scales on chin and throat
more elongate, distinctly keeled or tuberculated; scales on upper side
of fore limbs larger than dorsals, juxtaposed or imbricate, keeled, be-
coming larger and mnulticarinate toward the hand; scales on upper side
of hind limb similar, though less sharply keeled; scales on under side of
femur slightly larger than ventrals, indistinctly tuberculate; digital
expansion well developed, about thirty-three lamellse under second and
third phalanges of the fourth toe; tail strongly compressed, basal half
with a high fin-like crest supported by about fourteen bony "rays," the








SCI(11MJDT, AMI'HIBIANS OP PORTO RICO


elongations of the neural spines of the caudal vertebne; scales covering
sides of tail flat, keeled, those on the tin between the "rays" elongate,
three to four rows between rays, about fourteen longitudinal rows on
side of tail at the level of the fifth ray from the base: gular appendage
very large, with distant rows of small tubercnulate scales on the naked
skin, the edge being rounded, thickened, and scaly; large postanal
plates."
Female specimens lack the caudal "fin" and have a smaller dewlap
with the scales set more closely. Stejneger's description of the colora-
tion in life is as follows:
"Iris hazel, with a bright brassy ring bordering the pupil: general
color above greenish gray: back clouded with brownish and sides with
blackish dots, the dusky of the back and the black spots on the sides ar-
ranged in four perceptible, though indistinct, cross-bands; eyelids black-
ish, with a citron-yellowish spot above and behind the eye and a smaller
one in front; under the eye a long semilunar white spot barely invading
the posterior supralabials; several whitish spots on temples and sides
of neck; underside white with dark-gray mottlings and spots: dewlap
delicately Naples-yellow, scales on the edge white; legs indistinctly
crossbarred with dusky bands more or less spotted with blackish. Tongue
pale cadmium orange, whole interior of mouth of same color, but duller."
Stejneger records only a single specimen as being emerald green,
while ten of the eleven collected by me were a uniform green. The
eleventh was gray mottled with brown and black as described above.
It lay closely pressed to a small branch of which it seemed an integral
part until the continued pointing of my small boy assistant enabled me
to distinguish it. As a protective coloration this pattern is an extraor-
dinary success. Color-change is evidently very complete in this species.
There is little variation in the Porto Rico Survey series. In A. M.
N. H. No. 13234 the tail crest is unusually high, fully as high as in
A. ricordii of IIispaniola, but the scale characters which distinguish
cuvieri from ricordii are perfectly constant.
The measurements of the largest and smallest specimens are as follows:

Parts measured A. M. N. 11. A. M. N. H.
No. 13236 No. 13138
Total length .......................................... 418 mm. 304 mm.
B ody ................................................. 135 10)
T all .................................................. 283 204
Length of head ............. ..................... 43 32
Breadth of he d................ .. ........... 2(; 19
A rm .................................................. 60 39
L eg .................................................. 100 73








SCIENTIFIC SURVEY OF PORTO RICO


HIabits.-At Aibonito the giant Anolis was found in the coffee planta-
tions, usually on the larger shade trees, but occasionally on the coffee
bushes. It was never seen lower than eight feet from the ground. Tak-
ing into consideration its size, it is fairly abundant, but it is difficult to
see and still more difficult to secure without shooting.
Seven out of'eight stomachs examined contained the remains of large
beetles; one, a large phasmid; one, remains of Ileteroptera: one, a mass
of skin of Anolis cuvieri (doubtless its own). The boys say that this
lizard eats berries and fruits, and in the coffee plantations it is accused
of eating coffee berries. It seems probable that vegetable matter forms
only a small proportion of its food, as in Anolis crislallus. Wolcott
found snails in stomach contents examined by him.

Anolis cristatellus Dnmeril and Bibron
Lagartija comun
Text Figs. 27 and 28
Anolls cristatellus Dum6ril and Bibron, 1837, Erp6t. G4n., Vol. IV. p. 143.-
Dum6ril, 1851, Cat. Method. Rept. Mus. Paris, Vol. I. 1). 58.-Reinhardt
and Luetken, 1863, Vid. Medal. Nature. Foren., Copenhagen, 1862, p.
249.-Bocourt, 1873, Miss. Sci. Mex., Zool. Rept.. Livr. 2, Pl. 14, Fig. 12.-
Peters, 1876, Monatsber. Akad. Wiss. Berlin, p. 706.-Gundlach, 1881,
Anales Soc. Espaii. Hist. Nat., Vol. X, p. 300.-Stahl, 1882, Fauna Puerto-
Rico, p. 69, p. 159.-Boulenger, 1885, Cat. Lizards Brit. Mus., Vol. II,
p. 26.-Garman, 1887, Bull. Essex Inst., Vol. XIX, p. 27.-Meerwarth, 1901,
Mitt. Naturh. Mus. Hamburg. Vol. XVIII, p. 21.-Steineger, 1904, Rept
U. S. Nation. Mus., 1902, p. 638, Figs. 92-97.-Barbour, 1914, Men. Mus.
Comp. Zool., Vol. XLIV, p. 274; 1917, Proc. Biol. Soc. Wash., Vol. XXX,
p. 99.-Fowler, 1918, Papers Dept. Marine Biol., Carnegie Inst., Vol. XII,
p. 10. Fig. 5.-Schmidt, 1920, Ann. N. Y. Acad. Sci., Vol. XXVIII, p.
186.-Smyth, 1920, Rev. Agric. Pto. Rico, Vol. IV, p. 18.-Wolcott, 1924,
Journ. Dept. Agric. Pto. Rico, Vol. VII, p. 27.-Danforth, 1925, Copeia,
No. 147, p. 77.-Schmidt, 1926, Publ. Field Mus. Nat. Hist., Zool., Vol. XII,
p. 155.
Xiphosurus cristatellus Cope, 1861, Proc. Acad. Nat. Sci. Phila., p. 208.
Anolis monensis Steineger, 1904, Ann. Rept. U. S. Nation. Mus., 1902, p. 646.
Figs. 98-101.-Barbour, 1914, Mem. Mus. Comp. Zool., Vol. XLIV, p. 273.

Type locality.-Martinique (erroneously).
Distribution.-This species may be found in all parts of Porto Rico
except the more .densely forested areas. Combining the localities in the
Porto Rico Survey collection with those in Stejneger's list and those
given by Walcott, Anolis cristatellus is recorded from Adjuntas, Aguas
Buenas, Aibonito, Afiasco, Arroyo, Bayamon, Boqueron. Caguas, Cata-




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