Contributions to the geology and paleontology of the Canal Zone, Panama, and geologically related areas in Central Ameri...

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Title:
Contributions to the geology and paleontology of the Canal Zone, Panama, and geologically related areas in Central America and the West Indies
Series Title:
Smithsonian Institution. United States National Museum. Bulletin 103
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612 p. : ill., plates ; 25 cm.
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English
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Howe, Marshall A
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Govt. Print. Off.
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Washington
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Paleobotany -- Panama   ( lcsh )
Paleontology -- Costa Rica   ( lcsh )
Geology -- Panama   ( lcsh )
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federal government publication   ( marcgt )
bibliography   ( marcgt )
non-fiction   ( marcgt )

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Also available in electronic format.
Statement of Responsibility:
Marshall A. Howe et al.

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University of Florida
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oclc - 04332905
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U N I V UR S I Y or FLORYDA L I B R A R Y

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SSMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM Bulletin 103





CONTRIBUTIONS TO THE GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE, PANAMA, AND

GEOLOGICALLY RELATED AREAS IN CENTRAL AMERICA AND THE WEST INDIES








PREPARED UNDER THE DIRECTION OF

THOMAS WAYLAND VAUGHAN
Custodian of Madreporaria, United States National Museum,
Geologit in Charge of Coastal Plain Investigation,
United States Geological Survey
















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WASHINGTON
GOVERNMENT PRINTING OFFICE 1919









ADVERTISEMENT.
The scientific publications of the United States National Museum consist of two series, the Proceedings and the Bulletins.
The Proceedings, the first volume of which was issued in 1878, are intended primarily as a medium for the publication of original, and usually brief, papers based on the collections of the National Museum, presenting newly acquired facts in zoology, geology, and anthropology, including descriptions of new forms of animals and revisions of hmited groups. One or two volumes are issued annually and distributed to libraries and scientific organizations. A limited number of copies of each paper, in pamphlet form, is distributed to specialists and others interested in the different subjects as soon as printed. The dates of publication are recorded in the table of contents of the volume.
The Bulletins, the first of which was issued in 1875, consist of a series of separate publications comprising chiefly monographs of large zoological groups and other general systematic treatises (occasionally in several volumes), faunal works, reports of expeditions, and catalogues of type-specimens, special collections, etc. The majority of the volumes are octavos, but a quarto size has been adopted in a few instances in which large plates were regarded as indispensable.
Since 1902 a series of octavo volumes containing papers relating to the botanical collections of the Museum, and known as the Contributions from the National Herbarium, has been published as bulletins.
The present work forms No. 103 of the Bulletin series.
WILLI" Di&C. RAVENEL,
Administrative Assistant to the Secretary, In charge of the United States National Museum.
WASHINGTON, D. C., September 15, 1919.











PREFACE.
Geologists generally recognize that knowledge of the geology of
Central America is essential to solving the problems of the geologic history of the Americas, and many of them have devoted as much thought and study to the region as their rather occasional opportunities for investigation permitted. Among the previous investigators T. A. Conrad, W. M. Gabb, J. W. Gregory, W. H. Dall, H. Douvill6, P. Lemome and R. Douvill6, M. Bertrand and Ph. Zfircher, R. T. Hill, and Ernest Howe should be mentioned. Since work on the Panama Canal was initiated by the United States Government, excluding the investigations associated with official duties, contributions have been made by Franz Toula, A. P. Brown and H. A.
Pilsbry, and W. B. Scott.
In 1911 the Isthmian Canal Commission attached to its staff Dr.
Donald F. MacDonald as commission geologist. In October and No ember, in 1911, I had the privilege of spending a full month in field work along the canal, largely as a guest of the Canal Commission, and I here wish to express to Maj. Gen. Goethals, then Col.
Goethals, my very hearty thanks for the facilities afforded me.
Doctor MacDonald and I, of course, worked together, and he left
nothing undone in making our efforts successful.
Doctor MacDonald and I both recognized the extraordinary opportunity for making a highly valuable contribution, not only to the geology of Central America, but also to the geologic history of the
continents to the north and south. As a result of our conferences, I suggested to the Director of the United States Geological Survey a plan for cooperation between the United States Geological Survey, the Smithsonian Institution, and the Canal Commission. He approved the suggestion and submitted it to the Secretary of the 1 Smithsonian Institution, who also gave his approval. As a result 1 of these preliminaries the following letter was prepared and sent to
the chairman of the Canal Commission:
FEBRUARY 26, 1912.
Col. GEORGE W. GOETHALS,
Chairman Isthmian Canal Commission,
Washington Office, Washington, D. C.
Si: As a thorough knowledge of the geology of the Panamic Isthnian region is
essential to a solution of fundamental problems of the geologic history of both North South America and of the adjacent oceanic basins; as the excavations for the Panama Canal and along the line of the relocated Panama Railroad offer opportunities duringn g the next few years never before realized and probably never again to be realized III
!121525






I V PREFACE.

for a geologic study of this region; as there is a scientific need for the extension of the geologic investigations beyond the Canal Zone to adjacent areas, and as these extended investigations, although they may not always bear directly on the problems of building the canal, will, by furnishing a basis for a wider knowledge of the geology of the area than can be obtained on the Canal Zone, be helpful in deciphering the local stratigraphy and structure of the rock formations cut by the canal, and will afford information on whether there are fuels, notably fuel oil, or other geologic products of economic value within reach of the canal:
The Smithsonian Institution and the United States Geological Survey desire to enter into cooperation with the Isthmian Canal Commission in making a study of the geology of the Canal Zone and extending the studies to adjacent regions so far as is feasible.
The following is; submitted to the Isthmian Canal Commission for its consideration: It is hoped and urged that the Canal Commission will continue in its service a commission geologist, and will provide facilities for his field work within the Canal Zone until the excavations for the canal for the Panama Railroad, and for any other projects that may require excavations have been completed and carefully studied. The Canal Commission is especially requested to permit the commission geologist to extend his examinations of the geologic formations and mineral resources beyond the Canal Zone, the salary of the geologist to be paid by the Canal Commission, and funds for his field expenses to be provided by the Smithsonian Institution. The commission geologist Will, of course, submit to the Canal Commission a report of such nature and scope as the commission may direct.
The United States Geological Survey will, without charge, cut rock sections for microscopic study, make chemical analyses, and furnish special reports on fossils and other collections made and submitted by the commission. The advice of the different specialists on the survey will be at the service of the commission whenever their advice may be desired.
After the completion of the field work and after the commission geologist has submitted his report to the Canal Commission, the Smithsonian Institution desires to publish comprehensive and detailed monographic accounts of the physiography, stratigraphic and structural geology, geologic history, geologic correlation, mineral resources (including coal, oil, and other fuels), petrography, and paleontology of the Canal Zone and of as much of the adjacent areas in the isthmian region as is feasible. The services of the most eminent authorities will be enlisted in the preparation of special memoirs on the various collections made and submitted. The endeavor will be, by full presentation of all obtainable information, to make the Canal Zone the geologic standard of comparison for Central America as well as for portions of North and South America. In these reports due credit will be given to the Isthmian Canal Commission for its participation in the investigations.
We hope that, this plan will meet with your approval and support.
Vlery re 4pectfully,
(Signed) CHARLES D. WALCOTT,
Secretary, Smithsonian Institution.
(Signed) GEo. OTIS SMITH,
Director, U. S. Geological Survey.
The p)rop~osed cooperation was approved by the chairman 'of the Can1al Commilssion. Doctor MacDonald remained with the commission until1 the excavations in connection with the canal were completed and he nmade explorations outside the Canal Zone, especially along Banauia River in Costa Rica, and in the Province of Los Santos (Amuero Peninisula) and from David northward to the volcano of Chiriqui, In Panama. He was also geologist for the Costa Rica1'animi Boundary Commission.





PREFACE. V
Doctor MacDonald's reports to the Canal Commission have been published in the annual reports of the chairman of the Canal Commission; and he is the author of a more lengthy paper entitled "Some engineering problems of the Canal Zone in their relation to geology and topography," published as Bulletin 86 of the United States Bureau of Mines. 1 Since the termination of his services for the Canal Commission he has completed a large report on the physiography, stratigraphic and structural geology, petrography, and economic geology of the Canal Zone. The transmission of this memoir for publication has been delayed because some of the paleontologic determinations were needed for interpreting the geologic history.
After the agreement to the proposed plan of cooperation, I took charge for the United States Geological Survey of the preparation of the special paleontologic reports, of the problems of geologic correlation, and of the coordination of the investigations with other work on the physiography, stratigraphy, paleontology, and geologic history in the southeastern United States and the West Indies. The paleontologic material was sorted according to groups, and the following specialists undertook monographic reports:
Dr. Marshall A. Howe, calcareous algae.
Prof. Edward W. Berry, higher plants.
Dr. Joseph A. Cushman, foraminifera.
Dr. T. Wayland Vaughan, madreporarian corals.
Dr. Robert T. Jackson, echinoids.
Dr. C. Wythe Cooke, mollusca.
Mr. F. Canu and Dr. R. S. Bassler, bryozoa.
Dr. Mary J. Rathbun, decapod crustacea.
Prof. H. A. Pilsbry, cirrepedia.
The few vertebrates obtained were identified by Mr. J. W. Gidley. All of the paleontologic reports are now complete except that on the mollusks. It was at first hoped that Dr. W. H. Dall would prepare the one on this group, but pressure of other work prevented him. Later Dr. C. Wythe Cooke, paleontologist of the United States Geological Survey, began a study of the collection of mollusks, but other duties have interfered with his prosecution of it. The recent papers by Toula 2 and by Brown and Pilsbrys have been used, and they are valuable, but they do not meet the needs of the present investigation, for the material described in them mostly represents one geologic formation, the Gatun formation, and the stratigraphic
I U. S. Bureau Mines Bull. 86, pp. 88,29 pls., 9 text figs., 1915. Toula, Franz, Eine jungtertifire Fauna von Gatun am Panama-Kanal, Geolog. Reichsanstalt Wien Jbhrb., vol. 58, pp. 673-760, pls. 25-28,15 text figs., 1909; Die jungtertidre Fauna von Gatun am PanamaKanal, Ibid., vol. 61, pp. 487-530, pls. 30,31,1911.
3 Brown, Amos P., and Pilsbry, Henry A., Fauna of the Gatun formation, Isthmus of Panama, Acad. Nat. Sci. Phila. Proc. for 1911, pp. 336-373, pls. 22-29, 3 text figs., 1911; Fauna of the Gatun formation, Isthmus of Panama, 11, Acad. Nat. Sel. Phila. Proc. for 1912, pp. 500-519, pls. 22-26, 5 texts figs., 1913.




VI PREFACE.

data are not sufficient. It is probable that three and perhaps four horizons will be discriminated within the Gatun formation. Other groups of organisms are adequate for correlation purposes in most or all of the other geologic formations, but for the Gatun formation the principal reliance must be placed on the mollusks. The collections of mollusks made by Doctor MacDonald and myself is very extensive, and the greatest possible care was taken in obtaining full information on the stratigraphic relations of the material. It' is hoped that a report .commensurate with the size and importance of the collection may not be much longer delayed.
The series of papers here presented comprises all of the paleontologic memoirs that have been completed. These are immediately followed by descriptions of the geologic exposures where collections of fossils were made, with summaries of the fossils according to their stratigraiphic occurrence, and a chapter on the geologic correlation of the fossiliferous formations, both with other American and with European formations. It is intended that Doctor MacDonald's comprehensive general report will be published soon after this series of memoirs has been issued.
The names of the geologic formations use *d in the paleontologic reports are the same as those employed by Doctor MacDonald in Bulletin 86 of the United States Bureau of Mines, to which reference is made on page v of this preface.
I wish to thank the officials of the Canal Commission, particularly Maj. Gen. Goethals, Director George Otis Smith, and Chief Geologist David White of the United States Geological Survey, and Dr. Charles D. Walcott, Secretary of the Smithsonian Institution, for the support they have given these investigations. To my colleagues outside the Geological Survey and United States National Museum, Dr. Marshall A. Howe, Prof. E. W. Berry, Dr. Robert T. Jackson, Mr. F. Canu, and Prof. H. A. Pilsbry, who has collaborated in this work, I am under deep obligations; and it is a pleasure to record my appreciation of the efforts of my official colleagues, Dr. D. F. Mao Donald D)r. Joseph A. Cushman, Dr. C.W ythe Cooke, Dr. R. S. Bassler, Dr. 'Mary J. R athbun, and Mr. J. W. Gidley, all of whom have labored harmoniously to bring a, large undertakings to a successful conclusion.
THOMAS WAYLAND VAUGHAN.












CONTENTS.'


Page.
0ON SOME FOSSIL AND RECENT LITHOTHAMNIEAE OF THE PANAMA CANAL ZONE.
By Marshall A. Howe .................................................. 1
Introduction .......................................................... 1
Descriptions of species ................................................ 2
Archaeolithothamnium episporum .................................. 2
Lithothamnium vaughanii ......................................... 6
isthmi ............................................ 8
Lithoporella melobesoides ....................................11
Explanation of plates ................................................. 11
Index. ...................................................... i
'THE FOSSIL HIGHER PLANTS FROM THE CANAL ZONE. By Edward W. Berry... 15
Introduction .......................................................... 15
Correlation................................................... 16
Botanical character .................................................. 18
Tertiary ecology ...................................................... 21
Flora of the Canal Zone .............................................. 22
Systematic paleobotany ............................................... 23
Descriptions of species ................................................ 23
Fern fragments of Acrostichum ................................... 23
Palm rays ....................................................... 24
Palmoxylon palmacites ........................................... 24
Ficus culebrensis ................................................ 26
Guatteria culebrensis ............................................. 27
Myristicophyllum panamense ..................................... 29
Taenioxylon multiradiatum ........................................ 30
Inga oligocaenica ....................... ........................ 32
Cassi culebrensis ................................................. 34
Hiraea oligocaenica .............................................. 35
Banisteria praenuntia ............................................. 35
Hieronymia lehmanni ......................................36
Schmidelia bejucensis ............................................. 37
Mespilodaphne culebrensis ........................................ 38
Calyptranthes gatunensis ......................................... 39
Melastomites miconioides ............ ....................... .40
Diospyros macdonaldi ............................................ 41
Rondeletia goldmani .............................................. 42
Rubiacites ixoreoides ............................................. 43
Explanation of plates ................................................. 44
Index ................................................................ i
THE SMALLER FOSSIL FORAMINIFERA OF THE PANAMA CANAL ZONE. By Joseph
Augustine Cushman ..................................................... 45
Introduction. ................................................ 45
List of material ...................................................... 45
'For the most part the papers in this volume have individual indexes following the plates at the end of thepaper, and the Explanation of plates at the end of each paper gives a full description of the plates.
VII




VIII CONTENTS.

THE SMALLER FOSSIL FORAMINIFERA OF THE PANAMA CANAL ZONE-Con. Page.
Descriptions of species............................................... 51
Textularia abbreviata........................................... 51
sagittula............................................. 51
agglutinans............................................ 52
laminata ............................................. 52
subagglutinans........................................ 52
carinata............................................... 53
panamensis.......................................... 53
Chrysalidina pulchella.......................................... 54
Bolivina cf. B. punctata.......................................... 54
aenariensis.............................................. 54
robusta................................................ 55
species................................................. 55
Bigenerina nodosaria.............................................. 56
Gaudryina flintii ................................................. 56
triangularis.......................................... 56
Clavulina parisiensis............................................. 57
communis............................................. 57
Virgulina squamosa.............................................. 58
Lagena striata, var. strumosa...................................... 58
Nodosaria communis............................................. 59
insecta ................................................ 59
raphanistrum .......................................... 59
species ................................................ 60
Cristellaria rotulata.............................................. 60
italica............................................... 61
protuberans ......................................... 61
vaughani............................................ 61
Uvigerina canariensis............................................ 62
canariensis, variety.................................... 63
pygmaea............................................. 63
tenuistriata........................................... 63
Siphogenerina raphanus, var. transversus ......................... 64
Globigerina bulloides............................................. 64
inflata......................................... 65
dubia............................................... 65
conglobata............................................ 66
sacculifera........................................... 68
aequilateralis......................................... 67
Orbulina universa................ ................................ 61
Discorbisobtusa................................................. 68
Truncatulina americana............................................ 68
pygmea............................................ 68
ungeriana........................................... 69
wuellerstorfi ........................................ 69
culebrensis ........................................... 70
Pulvinulina sagra................................................ 70
concentrica........................................... 71
menardii............................................. 71
Siphonia reticulata.............................................. 72
Nonionina depressula............................................ 72
scapha...................................................... 73
panamensia......................................... 74
anomnalina............................................ 74






CONTENTS. IX

THE SMALLER FOSSIL FORAMINIFERA OF THE PANAMA CANAL ZONE-Con.
Description of species-continued. Page.
Polystomella striato-punctata ................................. 74
sagra .............................................. 75
macella ............................................. 76
crispa ................................................ 76
craticulata .......................................... 77
species............................................... 77
Amphistegina lessonii .............................................. 77
Quinqueloculina seminulum ..................................78
contorta ......................................... 79
auberiana ........................................ 79
undosa ........................................... 79
bicornis ......................................... 80
panamensis ....................................... 80
Sigmoilina tenuis ................................................. 81
asperula ............................................... 81
Triloculina trigonula .......................................82
tricarinata ............................................. 82
bulbosa. ............................................... 83
projecta ............................................... 83
Biloculina bulloides .............................................. 84
Spiroloculina excavata ...................................... 84
Orbiculina adunca ................................................. 84
Explanation of plates ..........................................85
Index ..................................................... i
T z LARGER FOSSIL FORAMINIFERA OF THE PANAMA CANAL ZONE. By Joseph Augustine Cushman ..................................................... 89
Introduction ................................................ 89
List of species and their geologic occurrence ........................... 90
Description of species ................ .... ...................... 91
Lepidocyclina canellei ............................................ 91
chaperi ............................................. 92
vaughani .......................................... 93
macdonaldi ........................................ 94
panamensis ......................................... 94
duplicata.......................... 96
Heterosteginoides panamensis ..................................... 97
Orthophragmina minima .......................................... 97
Nummulites panamensis ........................................... 98
davidensis ........................................... 98
Orbitolites americana ....................................... 99
Explanation of plates ................................................. 99
Index ................................................................ i
FossI ECHBNi OP THE PA.NAMA CANAL ZONE AN) COSTA RICA. By Robert Tracy Jackson ..................................................................................... 103
Introduction ......... ..................................... 103
List of species and their geologic occurrence ............................ 103
Description of species ....................................... .......... 104
Clypeaster lanceolatus ............................................ 104
gatuni ................................................. 105
Encope annectans ................................................. 106
platytata .................................................. 108
megatrema ............................................... 110
Echinolampas semiorbis ........................................... 112





X CONTENTS.

FOSSIL ECHINI OF THE PANAMA CANAL ZONE AND COSTA RICA-Continued.
Description of species-continued. Page.
Schizaster armiger ................................................. 113
cristatus ................................................ 113
panamensis ................. .......................... 114
Description of plates .................................................. 115
Index .......................................................... i
BRYOZOA OF THE CANAL ZONE AND RELATED AREAS. By Ferdinand Canu
and Ray S. Bassler ..................................................... 117
Descriptions of species ................................................ 117
Ogivalina mutabilis ............................................... 117
Cupularia umbellata ............................................... 118
canariensis ..........................................119
Holoporella albirostris ............................................ 120
Stichoporina tuberosa............................................. 121
explanation of plate .................................................. 121
Index ................................................................ i
DECAPOD CRUSTACEANS FROM THE PANAMA REGION. By Mary J. Rathbun. 123
Introduction .......................................................... 123
Literature on Tertiary Decapods of Panama ............................. 124
List of stations from which material has been examined, arranged from the
earliest to the latest, with the species found at each ................... 124
Descriptions of species .................................................. 131
Family; genus, and species indeterminable .............................. 131
Macrobrachium, species ........................................... 131
Macrobrachium (?), species .................................... 132
Nephrops costatus. .......................................... 132
Nephrops, species ................................................. 133
Pachycheles latus ................................................. 134
Petrolisthes avitus .................................................. 134
Axius reticulatus ................ .................................. 135
Axius (?), species .................................................. 136
Callianassa ovalis .................................................. 1 37
lacunosa ............................................... 138
elongata ............................................... 139
scotti ................................................... 140
crassim ana .............................................. 141
m oinensis .............................................. 142
spinulosa ............................................... 143
tenuis ................................................. 144
quadrata ............................................... 145
toulai .................................................. 146
abbreviata ............................................. 147
hilli ................................................... 148
vaughani ............................................... 148
stridiens ................................................ 151
m agna ................................................. 151
crassa .................................................. 152
Callianas.s species ................................................ 152
CallianaHsa (?), species............................................. 153
Petrochirus bouvieri .........................................153
Goniochele (?) arnata .............................................. 154
tiepat s (hilieinsis ................................................ 155
Ilepatus, species ................................................... 155






CONTENTS. Xl

DECAPOD CRUSTACEANS FROM THE PANAMA REGION-Continued.
Descriptions of Species-Continued. page.
Calappa costaricana ............................................... 156
flammea ................................... ............... 157
zurcheri .................................................. 157
Calappella quadrispina ............................................ 157
Mursia macdonaldi. ........................................158
obscure ..................................................... 159
Mursilia ecristata ................................................... 160
Leucosilia jurminei .......................... % ....................... 161
bananensis .. ........................................... 161
Leucosiidae, genus and species indeterminable................... 162
Callinectes declivis................ 162
reticulatus ............................................. 163
species .................................................. 164
Arenaeus, species .................................................. 165
Euphylax callinectias ............................................ 165
fortis. ................................... .......167
Gatunia proavita ................................................... 168
Carpilius, species .................................................. 171
Heteractaea lunata ................................................ 171
Panopeus antepurpureus ............................................ 172
tridentatus ............................................... 172
species ................................................... 173
Eurytium crenulatum .............................................. 174
Euryplax culebrensis .............................................. 174
Thaumastoplax prima ............................................. 176
Cardisoma guanhumi .............................................. 177
Uca macrodactylus ................................................ 177
Brachyrhyncha, family, genus, and species indeterminable.......... 177 Brachyrhyncha, family, genus, and species indeterminable........... 178
Parthenope panamensis ............................................. 178
pleistocenica ........................................... 179
Explanation of plates ................................................... 179
Index ................................................................ i
CIRRIPEDIA FROM THE PANAMA CANAL ZONE. By Henry A. Pilsbry ........ 185
Balanus eburneus .................................................. 185
glyptopoma ............................................... 185
concavus rariseptatus ...................................... 186
(Hesperibalanus?), species ................................. 187
Lepas injudicata ................................................... 188
Explanation of plate .................................................. 188
FossiLCORALS FROM CENTRAL AMERICA, CUBA, AND PORTO Rico, WITH AN
ACCOUNT OF THE AMERICAN TERTIARY, PLEISTOCENE, AND RECENT CORAL
REEFS. By Thomas Wayland Vaughan ................................... 189
Introduction ......................... ............................. .. 189
Geologic correlation by means of fossil corals ............................ 190
Geologic history of the upper Eocene and later coral faunas of Central
America, the West Indies, and the eastern United States .............. 193
Eocene .......................................................... 193
Brito formation, Nicaragua ............................... 193
St. Bartholomew limestone .....................................193
Jackson formation and Ocala limestone.......................... 195
Concluding remarks on the Eocene ............................ 198





Xii CONTENTS.

FOSSIL CORALS FROM CENTRAL AMERICA, CUBA, AND PORTO Rico, ETC.-Con.
Geologic history of the upper Eocene and later coral faunas of Central
America, the West Indies, and the eastern United States-Continued. page.
Oligocene......................................... ............ 198
Lower Oligocene .......................................... 198
Middle Oligocene ...................................... 199
Antigua formation ................... .................. 199
Pepino formation of Porto Rico ......................... 203
Limestone above conglomerate near Guantanamo, Cuba.. ...- 204 Basal part of Chattahoochee formation in Georgia ............205
Coral limestone of Salt Mountain, Alabama........... 206
San Rafael formation of eastern Mexico .................... 206
Tonosi, Panama........ ................................ 207
Serr'o Colorado, A rube................................... 207
Concluding remarks on the middle Oligocene................ 207
Upper Oligocene........................................... 208
Culebra formation................. ................ 208
Emperador limestone................................... 208
Anguilla formation ..................... ................ 209
Cuban localities........................................ 210
Tampa formation of Florida ............................ 211
Concluding remarks on the upper Oligocene..... 211
Miocene.. .................................................. 212
Bowden marl ............................................ 212
Santo Domingo......................................... 213
Cuba............................. ...................... 218
Baracoa and Matanzas ................................. 218
La Cruz marl ......................................... 218
Florida................... ................. a........... 219
Alum Bluff formation....... a........................... 219
Middle and South Atlantic States......................... 220
Costa Rica.................................. a ........ 221
Panama ................................ a................ 221
Colombia................................................ 221
Concluding remarks on the Miocene ......................... 221
Pliocene ................................................... 222
Caloosahatchee marl, Florida .............................. 222
Limon, Costa Rica ......a.................................. 223
Carrizo Creek, California................................... 223
Pleistocene ................................. ............... 225
Summary of the stratigraphic and geographic distribution of the Tertiary and Pleistocene coral-faunas of Central America and the West
Indies.................................................... 226
Table of stratigraphic and geographic distribution of species ..........228
Conditionsq under which the West Indian, Central American, and Floridian
coral reefs9 have formed, and their bearing on theories of coral-reef forma-!
tiort........... -.......................................... 238
Definition of the term "coral reef ".............................. 238
Ecology of reef-forming corals.................................. 240
htypothe:4es of the formation of coral reefs .......................... 241
Test,; of coral-reef hypotlieses ................................ 246
Criteri fo reogizn shift in position of strand line.......... 246






CONTENTS. XIIII

FoSSIL CORALS FROM CENTRAL AMERICA, CUBA, AND PORTO RICO, BTC.Continued.
Conditions under which the West Indian, Central American, and Floridian coral reefs have formed, and their beaxing on theories of coral-reef
formation. -Continued. Page.
Criteria for measuring the amount of vertical shift in strand line
and for determining the relative ages of terraces and the physiographic stage attained by a shore line ......................... 247
Criteria for ascertaining the r6le of corals as constructional agents. 248 Solubility of calcium carbonate in seawater..'. ............ 250
Effects of wind-induced and other currents in shaping coral reefs. 251
Criteria for determining the effect of glaciation and deglaciation
on the development of li i reefs ............................ 252
Amount of vertical displacement of strand line by glaciation
and degglaciation ......................................... 252
Rate of growth of corals and length of post-Glacial time ...... 253
Effect of lowering of marine temperature on reef corals during
glaciation .............................................. 254
Valley-in-valley arrangement and cliffed spurs .............. 256
American Tertiary and Pleistocene reef corals and coral reefs. . .. . 258
Eocene reef corals of St. Bartholomew .......................... 259
West Indian middle Oligocene reefs ............................. 259
Antigua .................................................. 259
Porto Rico ............................................... 260
Cuba .................................................... 261
West Indian and Panamanian upper Oligocene reefs ............. 262
Anguilla. .................................. 262
Canal 262
West Indian Miocene reef corals ................................ 263
West Indian Pleistocene reefs .................................. 263
Tertiary and Pleistocene reef corals and coral reefs of the United
States ..................................................... 265
Southeastern United States ................................ 265
Pliocene reef corals from Carrizo, Creek, California ........... 271
Living coral reefs of the West Indies, Florida, and Central America ... 271
Antigua Barbuda Bank ....................................... 273
St. Martin Plateau ............................................. 275
St. Croix Island ............................................... 278
Virgin Bank ................................................. 279
Cuba ........................................................ 280
Bahamas ...................................................... 291
Bermudas ..................................................... 293
Florida ...................................................... 297
Campeche Bank .............................................. 299
Honduran reefs ................................................ 300
Mosquito Bank ................................................ 300
Some other West Indian Islands ............................... 301
Brazil and Argentina .......................................... 301
Atlantic coast of the U nited States north of Florida ............. 303
Types of West Indian aud Central American littoral and sublittoral profiles and their relations to coral reefs ................ 303
Submerged banks north of the coral reef zone in the western Atlantic Ocean ................................................... 305
Summary of the conditions under which the American fossils and
living reefs formed .......................................... 305





XIV CONTENTS.

FOSSIL CORALS FROM CENTRAL AMERICA, CUBA, AND PORTO RICO, ETC.Continued
Conditions under which the West Indian, Central American, and Floridian coral reefs have formed, and their bearing on theories of coral-reef
formation .-Continued. Page.
Coral reefs of the Pacific Ocean ..................................... 306
Great Barrier Reef of Australia................................. 306
New Caledonia ................................................. 308
Fiji Islands .................................................... 309
Society Islands ............................................... 311
Tahiti ..................................................... 311
Smaller islands of the Society group........................... 312,
Atolls ............................................................ 313
Conclusions ....................................................... 319
Bearing of these conclusions on hypotheses of the formation of coral
reefs ........................................................... 325
Suggestions as to future investigations ............................... 329
Systematic account of the faunas ....................................... 333
Class Anthozoa ..................................................... 333
Madreporaria Imperforata ...................................... 333
Family Seriatoporidae ...................................... 333
Genus Stylophora ...................................... 333
Pocillopora ....................................... 342
Madracis ......................................... 345
Family Astrocoenidae ..................................... 345
Genus Astrocoenia .................................... 345
Stylocenia ......................................... 351
Family Oculinidae ........................................ 352
Genus Oculina ......... ................................ 352
Archolielia ........................................ 352
Family Eusmiliidae......... ..354
Genus Asterosmilia .................................... 354
Stephanocoenia .................................... 356
Dichocoenia ...................................... 360
Eusmilia .......................................... 361
Family Astrangiidae ....................................... 361
Genus Cladocora ...................................... 361
Family Orbicellidae ....................................... 362
Genus Orbicella ........................................ 362
Solenastrea ....................................... 395
Antiguastrea ....................................... 401
Stylangia .......................................... 410
Sepastrea .......................................... 411
Family Faviidae .......................................... 412
Genus Favia ........................................... 412
Favites., ......................................... 414
Goniastrea ......................................... 416
Maeandra ..........................................417
Leptoria ................. ........................ 421
Manicina .......................................... 421
Thysanus ......................................... 423
Family Mumsidae .......................................... 424
Genus Syzygophyllia ................................... 424






GONTENTS. XV

FOSSIL CORALS FROM CENTRAL AMERICA, CUBA, AND PORTO Rico, ETC. Continued.
Systematic account of the faunas-Continued.
Class Anthozoa-Continued. Page.
Madreporaria Fungida ........................................ 425
Family Agariciidae ....................................... 425
Genus Trochoseris.. ..............................425
Agaricia .......................................... 426
Pavona .......................................... 430
Leptoseris ....................................... 431
Pironastraea. --.................................. 432
Siderastrea ...................................... 435
Family Oulastreidae ...................................... 453
Genus Cyathomorpha ................................ 454
Diploastrea ....... ................................. 469
Madreporaria Perforata ....................................... 479
Family Eupsanimiidae ..................................... 479
Genus Balanophyllia ................................... 479
Family Acroporidae ....................................... 479
Genus Acropora ....................................... 479
Astreopora ....................................... 483
Actinacis ........................................ 486
Family Poritidae ........................................ 488
Genus Goniopora............................ ......488
Porites ........................................... 498
Class Hydrozoa .................................................. 507
Order llydrocorallinae .......................................... 507
Family Milleporidae ...................................... 507
Genus Millepora ..................................507
Explanation of plates ................................................507
Index ................................................................ i
THE SEDIMENTARY FORMATIONS OF THE PANAMA CANAL ZONE, WITH SPECIAL REFERENCE TO THE STRATIGRAPHIC RELATIONS OF THE FOSSILIFEROUS BEDS. By Donald Francis MacDonald ............................................ 525
Introduction .......................................................... 525
Sedimentary formations. ... .....................................526
Eocene (?) ......................................................... 526
Bas Obispo formation ........................................... 526
Las Cascadas agglomerate .................................. 526
Oligocene ....................................................... 526
Bohio conglomerate. .............................. ......526
Culebra formation .............................................. 527
Cucuracha formation. ....................................527
Emperador limestone ......................................... 531
Caimito formation ............................................. 531
Miocene ........................................................ 531
Gatun formation ........................................531
Panama formation .............................................. 532
Pliocene .......................................................... 532
Toro limestone............................................ 532
Chagres sandstone ............................................. 532
Pleistocene ........... --......................................... 532
Descriptions of local sections across the Isthmus of Panama ..............532
Section in canal cut 600 feet south of Miraflores Locks ................. 533
Section at Canal Commission station 2089 south of Miraflores Locks... 533
8370-19-11





XVI CONTENTS.

THE SEDIMENTARY FORMATIONS OF THE PANAMA CANAL ZONE-Continued.
Descriptions of local sections across the Isthmus of Panama-Continued. Page.
Section, north end of Miraflores Locks ............................. 534
Section, Pedro Miguel Locks to Paraiso Bridge...................... 534
Section at Bald Hill near Miraflores Locks........................... 534
Section along east side of Gaillard Cut from Canal Commission stations
1843 to 1850.................................................... 535
Section on west side of Gaillard Cut from Canal Commission stations
1775 to 1756.................................................... 535
Section on west side of Canal Commission station 1720, near Empire, to
1740, near Culebra.............................................. 536
Section on west side of Gaillard Cut near Las Cascadas, Canal Commission stations 1617 to 1597........................................ 537
Sections in cuttings of Panama Railroad near Caimito Junction ..... 539
Railroad cut near stream about midway between Rio Frijol and Rio
Frijolito....................................................... 539
Section in railway cuts near New Frijoles .......................... 540
Section showing chief railway cuttings and outcrops along the Panama
Railroad between Bohio and Monte Lirio......................... 540
Exposure a quarter of a mile northwest of old Bohio railroad station.. 541
Exposure opposite old Bohio railroad station, north side of the railroad
track........................................................... 541
Section at Pefia Blanca, about one mile below Bohio, on the west side
of Chagres River............................................... 541
Section at Yamos A Vamos, 21 miles below Bohio, west side of Chagres
River.......................................................... 542
Section on Panama Railroad from Monte Lirio to outcrop of Gatun
formation on south side of Big Swamp ........................... 542
Section showing Gatun formation, one-quarter to one-half mile from
Camp Cotton, toward Monte Lirio, at big curve on railroad........ 542
Large railway cutting a quarter of a mile from Camp Cotton, toward
Monte Lirio..................................................... 543
In the next two exposures going toward Camp Cotton ................ 543
Section in cut one-half mile west of Camp Cotton toward Gatun...... 543
Generalized sections of the bluffs exposed along the Panama Railroad,
relocated line, about 3,500 feet south of Gatun railroad station..... 543
Section from top of hill at western end of Gatun dam to bottom of the
spillway........................................................ 543
Section at west end of the spillway................................ 543
Exposures in the vicinity of Mindi Hill............................ 544
Monkey Hill, Mount Hope station.................................. 544
Section of )luff at end of Toro Point............................... 544
Section omne-third mile south of southern end of Toro Point Breakwater,
ini quarry....................................................... 545
THE BIOLOGIC CIIARACTER AND GEOLOGIC CORRELATION OF THE SEDIMENTARY
FORMATIONS OF PANAMA IN THEIR RELATION TO THE GEOLOGIC HISTORY OF
CENTRAL AMERICA AND THE WEST INDIES. By Thomas Wayland Vaughan. 547
Introduction........................................................ 547
Biologic character of the sedimentary formations in Panama.............. 547
Eocene ......................................................... 547
Oligocene........................................................ 549
lohio conglomerate ............................................ 549
Limestone on lIaut Chagres .................................. 549
Limestone at D)avid........................................... 549
Large Foraminifera from David ............................ 549





CONTENTS. XviI

THE ]BIOLOGIC CHARACTER AND1 GEOLOGIC CORRELATION OF THE SEDIMENTARY FORMATIONS or, PANAMA, ETC.-Continlued. Biologic character of the sedimentary formations in Panama-Con.
Oligocene-Continued. Page.
Culebra, formation........................................ 550
Fossils from the Culebra formation ....................... 551
Deposits of the age of the Culebra formatIon near Tonosi ......... 554
Larger Foraminifera from near To-nosi ..................... 555
Fossil corals from station ,6587, Tonosi..................... 555
Cucuracha formation ...................................... 555
Emperador limestone...................................... 556
Fossils from the Emperador limestone....................--557
Caimito formation........................................ 558
Miocene .................................................... 558
Gatun formation .......................................... 558
Fossils, except Mollusca, from the Gatun formation .......... 559
Mollusca from the Gatun formation, according to Brown and
Pilsbry............................................ 560
Pliocene.................................................... 562
Toro limestone........................................... 562
Pleistocene ................................................. 563
Fossils from the Pleistocene of the Canal Zone .................. 563
Correlation of the sedimentary formations of Panama................... 565
Tertiary formations of the southeastern United States ............ 565
A provisional correlation table of the Tertiary formations of the
South Atlantic and eastern Gulf Coastal Plain of the United
States ................................................ 569
Correlation of the Tertiary formations of the southeastern United
States with European subdivisions of the Tertiary................ 569
Eocene.................................................. 569
Oligocene................................................ 570
Miocene................................................. 572
Alum Bluff formation.................................. 572
Marks Head marl and the Calvert formation ................ 574
Choptank and St. Marys formations ...................... 575
Yorktown formation and Duplin marl..................... 575
Choctawhatchee marl.................................. 576
Pliocene................................................ 576
Age of the sedimentary formations of Panama, and the distribution of
their age-equivalents in Central America and the West Indies .. 577
Eocene.................................................. 577
Oligocene ............................................... 578
Lower Oligocene ..................................... 578
Middle Oligocene...................................... 582
Upper Oligocene ........................... .......... 585
Miocene................................................. 586
Pliocene ................................................ 593
Tentative correlation table of the Tertiary marine sedimentary
formations of Panama.................................... 595
Pre-Tertiary formations in CentralI America and the West Indies ...... 595





CONTENTS. Xviii

THE BIOLOGIC CHARACTER AND GEOLOGIC CORRELATION OF THE SEDIMENTARY FORMATIONS OF PANAMA,' ETC.-Continued. Outline of the geologic history of the perimeters of the Gulf of Mexico and
the Caribbean Sea ................................................. 596
Geographic relations of the three Americas ............ .............. 597
General relations ............................................. 598
Tectonic Provinces ............................................ 599
Baham as .................................................. 599
Atlantic and Gulf Coastal Plain ---------------------------- 60a
Mexican Plateau .......................................... 600
Oaxaca-Guerrero ----------------------------------------- 600Yucatan .................................................. 601
Guatemala-Chiapas ..................... ; ................... 601
Cuba ---------------------------------------------------- 601
Haiti, northern part ................................... 601
Honduras and, the Jamaican Ridge .......................... 602
Haiti, southerR part, Porto Rico, and the Virgin Islands 602 Saint Croix .......................................... 603
Costa Rica-Panama ....................................... 603
Andes ................................................... 603
Maritime Andes ------------------------------------------- 604
Caribbean Islands ---------------------------------------- 604
Barbadian Ridge ...................................... 604
Caribbean Are ---------------------------------------- 604
Aves Ridge ..................................... ..... 604
Paleogeographic summary .......................................... 604
Late Paleozoic ................................................ 605
Triassic, Jurassic, and Cretaceous .............................. 606
Eocene and Oligocene ----------------------------------------- 607
M iocene ------------------------------------------------------ 607
Pliocene and later ............................................ 609
Tabular summary of some of the important events in the geologic
history of the West Indies and Central America ............... 611






SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM Bulletin 103


CONTRIBUTIONS TO THE GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE, PANAMA, AND GEOLOGICALLY RELATED AREAS IN CENTRAL AMERICA AND THE WEST INDIES



ON SOME FOSSIL AND

RECENT LITHOTHAMNIEAE OF TH F PANAMA CANAL ZONE


By MARSHALL A. HOWE Of the New York Botanical Garden





Extract from Bulletin 103, pages 1-13, with Plates 1-11







~1.
PiR OR







WASHINGTON
GOVERNMENT PRINTING OFFICE 1918















ON SOME FOSSIL AND RECENT LITHOTHAMNIEAE OF
THE PANAMA CANAL ZONE.


By MARSHALL A. HOWE,
Of The New York BotaoicaIl Garden.

INTRODUCTION.
The following report is based chiefly upon a number of specimens of fossil calcareous algae, of the group known to geologists as" Nullipores," from Oligocene and Pleistocene strata in the Panama Canal Zone, collected in 1911 by D. F. MacDonald and T. W. Vaughan, of the United States Geological Survey.
In this material the Pleistocene period is represented by a single collection (MacDonald, 6039), consisting of numerous excellent free specimens, "from flats near Mount Hope, five feet above tide level." These Pleistocene specimens appear to the writer to belong to a species found by him a year or two earlier to be living in the Colon region, only a few kilometers distant. This species, so far as the writer can determine, has been hitherto undescribed; in framing its diagnosis, as published below, the fossil as well as the recent material has been considered, but a recent specimen, being more complete and satisfactory for detailed study, has been named as the technical type of the species.
So far as the present writer has been able to discover, the fossil coralline algae of America, in their taxonomic aspects at least, offer a practically untouched field for research. It is, of course, possible that geological and paleontological papers in which calcareous algae have been described have escaped the attention of phycologists, but inquiry among American geologists and paleontologists and a search of accessible literature have thus far revealed to the writer but a single' hitherto described species of fossil Lithothamnieae from the 'Western Hemisphere, namely, Lithothamnium curasavicum K. Martin, from the Island of Curacao, a species to which further allusion is made below in the discussion of Archaeolithothamnium episporum.
1Stromatopora compacta Billings (Palaeozoic Fossils, vol. 1, p. 55, 1862) from the Island of Montreal, etc., has sometimes been considered by geologists to be of corallinaus affinities (the species has been referred to Solenopora by Nicholson and Etheridge, 1Geol. Mag., vol. 3, p. 529, 1885), but, if we may judge from published figures, the organ.ism seems to the writer hardly a coralline alga, if indeed it is an alga at all.






2 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

The fossil Lithothamnieae of Europe have been described and figured in considerable number and with various degrees of care and detail. Most of these European descriptions and figures the writer has been able to see; some of them offer a reasonable basis for the future recognition of the forms concerned, without a reexamination of the original materials, but many of them do not. The present writer has had access to a good representation of the living Lithothamnieae of North America, the West Indies, Europe, and the East Indies, but so far as the fossil forms are concerned, he has had to depend upon descriptions and figures alone, which, as stated above, are often very unsatisfactory. In venturing to propose as new, two species of Lithothamnieae from Oligocene strata of the Panama Canal Zone, he doubtless risks the possibility that some future investigator, working with better materials or even with the same, may be able to convince himself or even to prove conclusively, that one or both of said species should be considered identical with species previously described from Europe. The diagnostic characters, the limits of variation, and the geographic range of even the living species are still very imperfectly understood. Some of the species are evidently widely distributed within certain temperature limits; others are at present known from single localities. So far as may be inferred from our present knowledge, very few, if any, of the forms of Lithothamnieae now living in tropical America occur also in European waters.

LIST OF SPECIES AND THEIR GEOLOGIC OCCURRENCE.
A.4rcihaeolithothamnium episporumin, new species. Recent, Toro Point; and
Pleistocene, Mount Hope; both in the Canal Zone.
Lithothamniunm vaughanii, new species, Oligocene, Culebra formation at
station 6026, about haif way between Monte Lirio and Bohio Ridge.
Lithothamniium isthmni, new species, Oligocene, Emperador limestone at
stations 6021, about 4 miles north of Gamboa Bridge, and 6024-b, Rio
Agua Salud, Panama Railroad (relocated line).
Lithlioporella melobesioides (Foslie) Foslie, Oligocene, Emperador limestone at station 6024-c, Rio Agua Salud, Panama Railroad (relocated
line).

ARCHAEOLITHOTHAMNIUM EPISPORUM, new species.
Plates 1 to 6.
Brownish red when living, the thallus forming at first widely epanded crusts 0.25-1.0 mm. thick, these in many cases repeatedly overgrown, the resulting crusts becoming 5 mm. or more thick, sometimes remaining nearly smooth or exhibiting the irregularities of the
SWe follow Rothpletz's original spelling of the final syllable of this unfortunately long name, a spelling that, happily, agreer with Philippi's spelling of the final syllable of LAthothamnium.




GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 3

substratum alone, but more often developing coarse, irregular rounded excrescences 5-12 mn. in diameter, or short rounded verrucae or nodules 2-5 mm. in diameter, the surface in sterile parts mostly smooth, indurated, and occasionally subnitent; hypothallia varying from weakly to strongly developed, 30-170 p. thick, their cells 17-28 V. by 8-11 p.; cells of the perithallium in distinct and regular layers except in oldest and youngest parts, the layers in more or less distinct zones, layers of short and of long cells occasionally alternating, cells mostly 8-15 IL by 5-8 p., in decalcified condition submoniliate, sphaeroidal to ellipsoidal, 1-2- times as high as broad, in calcified condition mostly subquadrate or oblong in vertical section: sporangia superficial, their apicula even with the surface, or slightly protruding, their cavities becoming only imperfectly and irregularly embedded, the sori slightly elevated, very irregular in outline, mostly 0.1-1.0 mm. broad, often widely confluent and anastomosing and becoming 5 mm. or more broad, the surface at length whitish and scarious, the ostioles mostly 16-22 p. in diameter, sporangia 65-96 p. high (including apiculum), 27-50 p. broad, 4-partite (occasionally 2-partite?), the spores irregularly paired or rarely subzonate.
Localities and geologic occurrence.-Covering dead corals, etc., and often forming concfetionary pebbles with coral cores, from lowwater mark to a depth of several meters, Point Toro, near Colon, Panama Canal Zone, Howe 6832 (type, in Herb. N. Y. Bot. Gard.), January 7, 1910; Colon, Howe 6840 (this covers continuously a mass of old coral 32 cm. long and 14 cm. in greatest width); also, as a Pleistocene fossil, "from flats near Mount Hope, five feet above tide level," D. F. MacDonald, station 6039,1 1911.
Paratypes.-Cat. No. 35298, U.S.N.M.
In outward form and in its habit of overgrowing old corals, Archaeolithothamniurn episporum resembles A. erythraeum (Rothpletz) Foslie, f. durum (Heydrich) Foslie, from the Red Sea and the East Indies, especially as illustrated by Weber-van Bosse and Foslie (Corallinaceae of the Siboga Expedition, pl. 5). Of this species we have seen only one specimen (from near Makassar), communicated by Mme. Weber-van Bosse, but from this and from the descriptions and figures of A. eiythraeumr published by Foslie, Heydrich, and Lemoine, we infer that the Panamanian specimens represent a different species. Perhaps the most important distinctive character of A. episporulm is to be found in its more superficial sporangia, as may be seen by comparing our photographs (pl. 2, fig. 1; pl. 3) with Heydrich's figure 2 of a vertical section through a sporangial sorus of his This is associated with minor amounts of other crustaceous corallines, including Lithophyllum, species, and Goniolithon, species. 2Ber. Dents. Bot. Ges., vol. 15, p. 68, fig. 2. 1897.





4 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

Sporolithon ptychoides, which Foslie I and Lemoine 2 consider to be synonymous with A. erythraeum. The sori or the emptied sporangial cavities appear also to be much less regularly embedded or overgrown by new tissue than is the case in A. erythraeum, if one may judge from Rothpletz's original description,3 Heydrich's figure 3,4 Lemoine's figure 29,2 and the descriptions given by the last-named writers; however, Foslie a remarks of .1. erythraeum that the sori are partly to be found overgrown in great numbers by new formed tissue, partly, however, they are not to be seen in section." In A. episporum, the sporangia themselves have never been seen except close to the surface; the emptied sporangial cavities do not show in a rough fracture or in an ordinary ground section, but irregular traces of them are often to be found in thin microtome sections of decalcified material. The sori of A. episporum are so superficial that their covering, after the discharge of the spores, appears to die and is flaked off together with more or less of the intersporangial parts, and the new tissue growing up from the base of the sorus shows only occasionatty and imperfectly the outline of the former sporangial cavities.
Rothpletz's original description of his Lithothamnium erythraeum leaves one in some doubt as to whether hlie found the contents of the sporangium divided or undivided; he uses the term Tetrasporen," but the measurements that he gives for these Tetrasporen are such as commonly belong to the whole sporangiumi in this group. In Heydrich's first description "' of his K.porolithon ptycwhoides, the "Tetrasporangien" are said to be meist ungetheilt, selten zweitheilig," but a little later hlie figures four tetrapores in a sporangium, arranged in the cruciate" manner. But this mode of division being at variance with the prevailing ideas as to the arrangement of the spores in the Corallin'ae. F'oslie. a little later in writing a diagnosis of the genus i rchaeolthothamniuw. inserted a question mark after "sporangia * uiparted or cruciate?" and this sign of doubt as to the cruciate division has been repeated by later writers. In A. episporum the mature sporangia are commonly and normally 4-parted in an irregularly cruciate fashion, but often the division axes of the two pairs of spores are at right angles to each other, so that only three spores are visible in a lateral view, and occasionally
S ibofar Exped. Monog., No. 61, p. 38. 1904.
Ann. Inst. Ocdanog., vol. 2, pt. 2, p. 67. 1911.
:lRothplotz, A. Bot. Centralb., vol. 54, p. 5. 1893.
S eor. ieuts. l ot. ( s.. vol. 15., p. S. 1897.
Nibog Expeld. Monog., No. 61, p. 41. 1904.
I Ber. l)euts. Bot. s vol. 15. p. 49 7
Sldern, pl. 18, fig. ;..
Kgl. Norske Vidensk. Selsk. Skr. 1900, p't. 5, p. 8. 1000.
I)Do Toni. Syll Alg., vol. 4. p. 1 721, 1905 Svodelius, in Eng. & Pi'rantl. Nat. Pflanzenfurn vo|.1, pt. 2 N (cht-i ige,. ). 2;7. 1911.





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 5

the second divisions seem to be omitted and the sporangium is apparently mature with only two spores. Very irregular types of division also occur, and rarely one finds an approach to the zonate I arrangement characteristic of most of the Corallinaceae.
The perithallic cells of A. episporum appear to be, in the decalcified state, more rounded and in more moniliform filaments than is the case in A. erythraeum, as may be seen by comparing our photomicrograph 2 with the photomicograph of a presumably decalcified section of A. erythraeum-published by Lemoine.Y The distinct stratification of the perithallium of A. episporum is due, in part, to the alternation of layers of long and short cells, but we have never seen in the Panamanian species any such striking alternation of long and short cells as is shown in this photograph published by Mine. Lemoine and as is shown still more emphatically in Heydrich's figure 3 of a vertical section of his Sporolithon ptychoides.
From Archaeolithothamnium dimotum Foslie and Howe,5 the only living species of this genus previously described from the West Indian region, A. episporum differs widely in its thicker crusts, in its more superficial sporangial sori, which are for the most part exfoliated after maturity of the sporangia and are only obscurely and imperfectly overgrown, in the usually larger, more rounded, and more moniliately arranged cells of the perithallium, the larger and rather less widely separated sporangial ostioles, etc.
Archaeolithothamnium curasavicum (K. Martin) Foslie,6 a Cretaceous fossil from the island of Curagao, is described and figured as showing distinctly rows of embedded sporangial cavities, such as would not be seen even in a thin decalcified section of A. epi.sporumn.
A Pleistocene fossil, collected by MacDonald at station 6039, from flats near Mount Hope, came from a few kilometers from the localities where we found the plant living, and we can entertain no serious doubt as to the specific identity of the recent and the fossil forms. The living and fossil are similar in external habit, as may be seen by comparing plates 1 and 4. They are similar also in their relations to old corals, and in structure (compare fig. 1, pl. 2. and fig. 4, pl. 5) they appear to exhibit only such differences as may be ascribed to individual variation or as may be expected in comparing the recent or living with the long dead. But little remains of the fossil speci'Zonately 4-parted sporangia have been described by Foslie for the Californian Archaeolithothamnium zonatosporum (Foslie, Algologiske Notiser. II. Kgl. Norske Vidensk. Selsk. Skr., 1906, pt. 2, p. 14), so that it would appear that this genus exhibits a wide variety in the matter of division of its sporangia.
SPlate 3, fig. 2.
Ann. Inst. Ockanog., vol. 2, pt. 2, pl. 1, fig. 1. 1911.
4Ber. Deuts. Bot. Ges., vol. 15. 1897.
SBull. N. Y. Bot. Gard., vol. 4, p. 128, pl. 80, fig. 1; pt. 87, 1906.
6 Lithothamnium curasavicum K. Martin, Bericht iiber eine Reise nach Niederlindisch West-Indien und darauf gegrindete Studien. II. Geologie, p. 26, pl. 2, figs. 22-25, 1888.




6 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

mens after decalcification, though the outlines of the cells may be recognized here and there. As microtome sections of the decalcified fossil material are out of the question, comparisons of "tructure of the recent and fossil must naturally be based upon calcareous ground sections. And in comparing the cell structure in sections of the recent decalcified specimens (pl. 3) with that shown in ground sections of the calcareous fossils, it is necessary, of course, to bear in mind that cells in calcareous ground sections of the Corallinaceae comminonly appear much more rectangular than in decalcified sections of the same material.' In the sections of the fossil material thus far made there are no certainly recognizable traces of sporangial cavities, but this is true in almost an equal degree of calcareous ground sections of the recent specimens except as to the surface of the plant (fig. 1, pl. 2), where the sori are, in fact, so decidedly superficial or even exserted that they could, perhaps, hardly be expected to persist in the fossil state.
In the same locality with the type-specimens (Howe 6832) there occurs an outwardly somewhat similar plant (Howe 6837) that we at first suspected to be the antheridial form of A. episporum, but certain recognizable, though possibly unimportant, differences in the form, size, and zonation of the perithallic cells have restrained us from so considering it. The antheridial conceptacle (cavities) in this 6837 are 64-95 p. broad and 60-72 high; they become copiously embedded by the continued upward or outward growth of the thallus.
LITHOTHAMNIUM2 VAUGHANII, new species.
Plate 7. 1igs. I nd 2, and plate S.
Thallus forming at first expanded crusts 1-2 mm. thick, these becoming overgrown. irregularly stratified, and 10 rum. or more thick, developing finally numerous, rather carse, crowded anastomosing branches, and forming masses 2-4 cm. or more high; branches mostly 3-12 mm. in diameter, usually much flattened, occasionally subterete, often reduced to anastomosing ridges, or sometimes appearing as dome-shaped elevations 2 cm. or more broad; primary hypothallia somewhat reduced, their cells 14-33 IL by 8-14 ., rather irregularly arranged (i. e., not distinctly "coaxial "), cells of medullary hypot1 llia mostly 15-30 p by 5-13 ., secondary hypothallia numerous and thin; branches showing in section numerous narrow irregularly flexlious, often subelliptic-lenticular or subcrescentic zones caused by
IFor illustrations of these differences, see Lemoine, Ann. Inst. Ocanog., voJ. 2, pt. 2, p. 45, figs. 19-21, 1911.
'"hi. writer believes, with Mine. Paul Lemolne, that the current rules of nomenclature require that Philippi's original spelling of this generic name should be respected, even though prevailing usnge has modified the final syllable. Whether the rules of nomenclatuio justify the use of this generic name for any of the species now bearing it is a more comrplieated oust ion.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 7

the alternation of layers of short and long perithallic cells, or by the interpolation of reduced secondary hypothallia; the larger perithallic cells mostly 13-122 V. by 11-14 p, usually higher than broad, the smaller subquadrate, about 8 p. square, or sometimes much compressed (7. high, 14 p. broad); conceptacles becoming embedded; tetrasporic conceptacles much flattened, oblong or elliptic-oblong in radio-vertical section, the cavity 500-740 V. in maximum width, 130-230 p. in height; roof of the tetrasporic conceptacle rather sharply defined, its cells in regular vertical rows of 1-4 cells, often elongate vertically, becoming sometimes 25-30 V. high.
Locality and geologic occurrence.-Oligocene, Culebra formation, "about half way between Monte Lirio and Bohio Ridge, on the relocated line of the Panama Railroad," collected by D. F. MacDonald and T. W. Vaughan, 1911 (station No. 6026).
Holotype and paratypes.-Cat. Nos. 35299, 35300, U.S.N.M.
The specimens obtained are more or less embedded in a hard rock matrix, so that our photograph (fig. 1, pl. 7) can give only an imperfect idea of the outward form of the plant. With a little mental clearing away of the matrix, it seems probable that in size and ext'ernal appearance, the species may be compared with rather coarse eroded conditions of the living Lithothanmniu rm glaciale Kjellman, but there is little similarity in structure; the perithallic cells of L. vaughanii average considerably larger than those of L. glaciale and they are arranged in more distinct layers; the embedded tetrasporic conceptacles of L. vaughanii are more flattened than those of L. glaciale, their cavities have about twice the maximum width of those of L. glaciale and the specialized character of the conceptacle roof is not noticeable in L. glaciale.
In external habit Lithothamnium vaughanii may perhaps be compared also with the living Lithophyllum racemus (Lamarck) Foslie forma crassum (Philippi) Foslie' of the Mediterranean and Adriatic seas, especially as shown in Hauck's figure 2 2 under the name Lithothamnium crassum Philippi, though the Panamanian fossil sometimes develops longer and perhaps more flattened branches than this form.
Of the living Lithothamnieae now known to the present writer as occurring in the West Indian region, Lithothamnium vaughanii per1 Kgl. Norske Vidensk. Selsk. Skr., 1898, pt. 3, p. 9, 1898. Foslie's identification of Lithotham nium crassum Philippi as a form of Lithophyllum racemus (Lamarck) Foslie was accepted by Heydrich (Bot. Jahrb., vol. 28, p. 536, 1901), but Mme. Lemoine quotes Lithothamtum crassum Philippi as a synonym of Lithothamnium caleareum (Pallas) Areschoug. It is not, however, apparent that any of these writers examined authentic material of Philippi's Lithothamnium erassum, if such exists. It is of some interest, also, to note that less than six months before Heydrich accepted Lithothamnium crassum Philippi as a form of Lithophyllum racemus he named it as the type of a proposed new genus Stichospora (Ber. Deuts. Bot. Ges., vol. 18, p. 316, 1900).
Hauck, F. Die Meeresalgen Deutschlands und Oesterreichs. In Rabenhorst, L., Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz, vol. 2, pl. 1, 1885.





8 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

haps most resembles Lithophyllum daedaleum Foslie and Howe 1 as to general habit, but differs from it much in structure.
In the best section, No. 35299 U.S.N.M., the one from which the photographs (fig. 2, pl. 7 and pl. 8) were made, the coarse intersporangial sterile tissue of the tetrasporic conceptacles is scarcely shown, yet the roofs of the conceptacles show unmistakable canals and none of the conceptacles in section exhibits a single orifice, so that we consider ourselves justified in inferring that the specimen in question is tetrasporic and that it belongs in the genus Lithothamnium in the sense in which that name is currently applied to living plants. In a section from another specimen under the same collection number, traces of the sporangia and of the intersporangial sterile tissue are evident. It is to be observed also that the zonate arrangement of tissues, as observed in a section, is essentially of the character assumed by Mme. Lemoine 2 as being peculiar to the genus Lithothamnium. The rather distinctly specialized nature of the cells of the conceptacle roof is evidently a character of importance, in which respect it differs markedly from the plant we are describing as Lithothamnium isthmi, as also in the distinctly zonate structure of the thallus, the reduced hypothallium, the larger tetrasporic conceptacles, larger perithallic cells, etc.
Among the more fully described fossil Lithothamnieae, L. vaughanii may perhaps be compared with Lithothamnium suganum Rothpletz 3 from the Tertiary (" Scio-Schichten ") of Val Sugana, near Borgo in the Austrian Tyrol, but the conceptacles of the Panamanian fossil are much larger (500-740 p wide and 130-230 V. high vs. 250 pL wide and 100 p. high) and the perithallic cells appear to average considerably larger, being sometimes 13-22 p. high, while those of L. suganumr are described as 9-12 p. long.

LITHOTHAMNIUM ISTHMI, new species.

Plate 7, fig. 3; plates 9, 10, and 11.
Thallus forming at first stratified crusts 3-12 mm. thick, but at length developing tortuous anastomosing branches and forming large rather solid, concrescent, fruticose masses; branches mostly 2-12 mm. in diameter, much flattened or subterete, often subconic-cylindric, flexed-digitiform, or molariformn; hypothallia showing regular concentric layers of cells (" coaxial"); hypothallium of the crustaceous parts 160-480 p. thick, its cells 17-28 p. by 8-13 p, transition to the
SBull. N. Y. Bot. Gard., vol. 4, p. 133, pls. 83, 84, 93, 1906.
SLemolne, Mine. Paul. Structure anatomique des Mlobesles. Application a la classiication. Ann. Inst. Ocanog., vol. 2, pt. 2, pp. 27, 28, 1911. Zelts. Deuts. Geol. Ges., vol. 43, p. 319, pl. 17, fig. 4, 1891.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 9

perithallium abrupt; medullary hypothallium of the branches mostly 0.6-2.0 mm. in diameter, often turning yellow and more or less disintegrated, its cells 17-44 p. by 8-13 I, transition to the perithallium abrupt or gradual; cells of the perithallium in distinct layers, the layers in rather indistinct zones; perithallic cells of the crustaceous parts subquadrate, 8-11 in diameter, sometimes only 6 p broad; perithallic cells of the branches usually a little higher than broad, 8-19 p. by 8-12 .; conceptacles becoming embedded; tetrasporic conceptacles appearing much flattened in a vertical section, the cavity 240-550 p. in maximum width, 130-165 p. in height.
Localities and geologic occurrence.-In Emperador limestone of Oligocene age (and often constituting the dominant element in its composition) on relocated line of the Panama Railroad, opposite San Pablo, Panama Canal Zone (" first limestone outcrop just north of Caimito Station, about four miles north of Gamboa Bridge "), collected by D. F. MacDonald and T. W. Vaughan, 1911, Station No. 6021 (No. 35301, type); and "above foraminiferous marl at Agua Salud Bridge about -1 mile north of New Frijoles on relocated line, Panama Railroad," by the same collectors, Station No. 6024b.
Holotype and paratypes.-Cat. Nos. 35301 to 35303, U.S.N.M.
The material upon which the above description is based shows much variation in form and structure and it was our first impression that two or more species were represented in it. However, if this is true, the two or more species are so intergrown and entangled and are so similar in structure that it is difficult to determine where one begins and the other ends. As regards the vegetative structure, we believe that we have been able to trace the continuous organic connection of the two types shown in our photomicrographs (pl. 9 and fig. 2, pl. 11), yet it is notoriously easy in the case of overgrowing and overgrown fossil Lithothamnieae to mistake the close contact of independent plants for structural continuity.
In the tetrasporangial specimen (No. 35301-fig. 3, pl. 7 and pl. 9) that we have named as the type, the thallus presents itself in the form of irregularly superposed crusts, more or less overlaid by crusts showing a somewhat different structure and conceptacles of a different sort, these outer layers probably representing a crustaceous species of Lithophyllum. The hypothallium of this No. 35301 is suggestive of that figured by Foslie for his living Lithothamniumn fragilissimum from Borneo (which, however, has a much thinner thallus). It suggests also the hypothallium of Lithothamnium lich1 Foslie, M. Lithothamnioneae, Melobesieae, Mastophoreae. In Weber-van Bosse, A., and Foslie, M. The Corallinaceae of the Siboga Expedition, Siboga Exped. Monog. No. 61, fig. 5, 1904.






10 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

enoides, as figured by Rosanoff 1 and by Lenioine, 2 but the crusts are evidently more massive than in that species.
Although the outward form of Lithothimium, isthi is more or less obscured by bein g embedded in rock, it seems; probable that in its typical condition (No. 35301) the external appearance of the plant may be compared with the recent plant from the Adriatic Sea, figured by Hauck3 -is "Lithwphylam decwti .ahtw Solims," wNihich Foslie 4
afterwards referred to his Litho thammiuin philippi-a species that he maintained even after conceding5 its specific identity With the' earl ier-publ ished Lithophyltum, e ispatumn, Ha tick. The typical form of Foslie's L~hotham?,niutiia philhppii is said by him" "to have its hypothallium distinctly marked and vigorotisl 'y developed, forming a coaxilate layer," but the "coaxial" character is essentially denied by Mine. Lemoine -- to what she considers the same spIecies under' the name Litliotham hun ci-ispatI1 Hlauck. The perithallic cells of the crustaceotis pa~rts of Lithothamiimz ?'thm; ,1ppe)ar to average considerablv sin- 11cr thian tlios e of Li. (LP5hftr 1. phi/,p p;;) according to the measurements given byLmieand by 1Foslie. The tetrasporangaial conceptacles of the Lithoph?/llum decllsa?1iit of Hauck (Lithothamuitn philippiil Foslie) are stated by 1-muck to be "8Q0,. bis -1 imll In di meter, while in- Lit hot hain,m ;u ;sthrni they are only 240-550 V. in maximumiii width. Moreover',
unlsswe 1:ue Mistaken InI connectino-r the fiuticulose parts of the Panamau~nian-fos'ii with the crusts, Lithotlwmniurnt i'sthm!i develops win1UC170U5 solid .inastoinlosinig branches, while in 1. crsptm the
shortbranchlike excrescences are mostly hollow, infu-ndibuliform, 01 scphiforin. These fruticulose conditions, which comprise a larir-e part of the ma,,terial collected by MacDoinld and Yaughaitn s uggest. in external form certain states of the living West Indian L,;t,'hph~y7llum daedalcumt Foslie and H~owe, which also presents
itself in both crusta,_ceous and fruticulose conditions. Occasion-ally
anl unu11sually long subterete branch may resemblie in form a fra1 Mom. 1oc. im. 'Sc. Nait. Cherbourg, vol. 2,pl. 6, fig. 14, 180G.
SAnn. Iiist. Oc6:nog-., Vol. 2, pt. 2, fig. 60, 1911. It is of interest to note that Mmne. Lertnoinr. basing- her system of classification primarily upon thie Vegetative structure of t le al hi, cves LIAtho th(wmilium lich en oi(es in the genus Lithiop/i yllum, notwi thstand1ogth fact thlat, its tot rasporangia are borne as it) the genns Litli otltamiiniu of modern wrIer Hite sam11e way~ Ahe wou11l doub~tless place Lithothamntiurn ist/mi in thie genus, I,0ilohp hihn, evenl though th1is species (or- it', type at least) clearly hmas the totrasporang 'IaI conc]efpt acloes of thIIe CoI V cnt Iionial lit ho 11 il 11illlo.I
:: 1ick. F. lioM*rle Dcutschilanlds ud Gesterreichis. pl. 1, fig. 7. Set, also
()1. 1, fig". 1, of Foslie's Die Lithothainnien des, Adriatischen Mleeres und Mlarokkos (WISS.
Meeresuntersu h lolmd, Vol. 7, pt. 1, 1904).
Foslio, M. Oni some L ithothiamnla. Kgl. Noreke Vidensk. Selsk. Skr., 1897, pt. 1, j,7, 18R97T.
NVWISS. 'Meeresuntersuch, flelgoland, vol. 7, pt. 1, pp. 13, 14, -1904.
6Kgl. Norsko Yidensk. Selsk. Skr., 1900, pt. 1, p. 5, 1900.
Anti. ist. Oc(-anog., vol. 2, pt. 2, p. 80, fig. 28, 1911.




GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 1 1

ment of the living East Indian Lithothamnium pulchrum A. Weber and Foslie.1
Lithothamnium fosliei (Trabucco) De Toni (Syll. Alg., vol. 4, p. 1761, 1905), a Miocene fossil from Italy, is figured 2 as having a "Lcoaxial" hypothallium, but from the illustrations given of the conceptacles, there is no sufficient ground for considering this plant to be a Lithothamnium rather than a Lithophyllum. In the original place of publication nothing but a figure (section) is given, from which, according to the scale of magnification given, it would appear that the conceptacles are only 140-160 I by 80-90 a and the perithallic cells about 16 i high, making the cells rather larger and the conceptacles much smaller than in L. isthmi.
If we are correct in including with Lithothanium isthmi the more ramified forms collected by MacDonald and Vaughan, the species, though commonly coarser, appears to be sometimes suggestive of plants figured as Nullipora rarmosiseima Reuss or Lithothamnium ramosissimum (Reuss) Schimper, from the Tertiary "Leithakalk" of the vicinity of Vienna, but Reuss's original figures and description relate to external form only, and give no adequate basis for referring the plant to a modern genus. Unger' adds good figures of the vegetative structure, but shows no conceptacles. Rothpletz 5 describes the conceptacles of L. ramosissimum as 280 p high, while the height of the conceptacles of L. isthmi is 130-165 p. and the width 240-550 p.. Rothpletz has no doubt that there are two species of Lithothamnieae in the Leithakalk," which may have been confused.
LITHOPORELLA MELOBESOIDES (Foslie) Poslie.
Lithoporella melobesioid.es (Foslie) FOSLIE, Kgl. Norske Vidensk. Selsk.,
1909. pt. 2, p. 50.
Mastophora inelobisioidesc FOSLIE, Kgl. Norske Vidensk. Selsk. Aarsber.,
1902, p. 24, 1903.
Locality and geologic occurrence.-This species occurs in small quantity with Lithothamnium isthmni in Emperador limestone of the Oligocene age, "above foraminiferous marl at Agua Salud Bridge about -1 mile north of New Frijoles on relocated line, Panama Railroad,"D. F. MacDonald and T. W. Vaughan, 1911, Station No. 6024b.
EXPLANATION OF PLATES.
PLATE 1.
Archaeolithothamnium episporuin M. A. Howe.
Photograph, natural size, of the type-specimens. collected at Point Toro, near Colon, Panama Canal Zone, January 10, 1910 (Howe 6832). The technical Compare pl. 4, Siboga Exped. Monog. 61.
2Eulithothamniumn fosliei, Trab. Boll. Soc. Geol. Ital., vol. 19, pl. 11, fig. 10, 1900. aHaidinger, Naturw. Abh., vol. 2, pt. 1, p. 29, pl. 3, figs. 10, 11, 1848. Type from Nendorfl," Hungary.
SDenkschr. k. Akad. Wiss. Wien, vol. 14, p. 23, pl. 5, figs. 18-22, 1858.
3 Zeits. Deuts. Geol. Ges., vol. 43, p. 320, 1891.
8370*-18h-Bull. 103- 2






BULLETIN 1413, UNITED STATES NATION AL MUSEUM.

type in a narrower sense is the specimen shown it the lower right-hand corner of the plate--the specimen from which figure 1 of plate 5 w~as obtained.

PLATE 2.

A rCJchaoli t iiof ,)i~iiii ai epiI ) cnsporut M. A. Howe.

Photographs of radio-vertical ground (calciferous) sections of type material (Point Trhoio, Howe 6882).
Fics. 1. kithothaiiiijii(tu iqihu-Al A. H1owe. Photograph of the typeportion of ai sporangial sorus, enlarged 42 diameters.
2.~ ~~t Seton!nlred 200 diameters.

PLATE 3.

A reolith o? i(i O/i(iihi. p i l)ipor U? M. A. Howe.

P'liotograpi is of radio-vertical sections of dec ilci fled material (Point Toro, Howe 6832), enlarged 200 diameters. 11"w. 1. Sectioni showing sporangtia and tetraispores.
2. Section showing e-mptied sporangia, form and arrangement of penithallic cells, a weakly developed hypothallium, etc.

PLATE 4.

Ar-ch iolitUi 0tha (Din jiil11 episporurn M. A. Howe.

A Pleistocene fossil, from flats near MVount Hope, five feet above tide level," D F. MNacTonald 6039, 1911, natural size.

PLATE 5.

A rclai oli tlol hliit i? i urn ('jisporulIn Al A. Howe.

Fiecs. 1 and 2. Photographs of the type maitteria-l (Point Toro. Howve 6832). Fic.. 1. Portion of the surface, showing the more or less confluent sporangial
son, enlarged 4 diameters.
2A smaller part of the same surface, showing the spor.,ngial ostioles,
etc., enlargedl 25 diameters.
Views. 3 and 4. Photograiphs of Pleistocene specimien from Mount Hope (MaclDonald, Cat. No. 35298, U.S.N.M.)
Vie. 8. I~imv(enI ical section showing soeeral superposed cru'ists anid three welllevelopedl hypothall ia. enlarged 42 dliameters.
4. .\ part of n cross4, -wct ion of one of' the excrescencees or brantiches, showing
.I single wealkly developed hypothailliumn, en~largedl 42 diatmeters. Compa;re, 40 r1c01re 40, living- spec-imenl as shown in fig. 1, Jplate 2.,

PAE6.

I rh aold (Jllun flm pi.sporion~ M,. A. Howe.

Photog (ip owt (ion of thf lesocn fossil from niear M)ount Hope (CMt o 59,SNM A eilionmagified t73 diniiieter.s.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 13

PLATE 7.

Io. 1. Lithiothamnium vaughlianii M. A. Howe.. Photograph of the typespecimens (between Monte Lirio and Bohio Ridge, MacDonald and
'Vaughan. Cat. No. 35299. U.S.N.M.), natural size.
2. Lithothamnnitam vauglhanii. A section showing irregular zonation, tetrasporic conceptacles. etc., enlarged 42 diameters.
3. Lithothanmiumn isthmi M. A. Howe. A section, slightly enlarged (11/8
of the natural dimensions), showing the type-specimen embedded in the matrix (from about 4 miles north of Gamboa Bridge, MacDonald and Vaughan, station 6021). The type material (Cat. No. 35301, U.S.N.M.), from which the section shown in plate 9 was obtained, occupies the central portion of the light area and is overgrown by crusts of what appears to be a different plant. probably a species of
Lithioph yllum.

PLATE 8.

Lithothaamnium rauihianii M. A. Howe.

An enlargement of a. part of the section shown in figure 2, plate 7, illustrating form of perithallic cells, the reduced secondary hypothallia, the somewhat specialized roof of the tetrasporic conceptacles, etc. Magnification 100 diameters.

PLATE 9.

Litliotihamniumn isthmi M. A. Howe.

A section of the type material (MacDonald and Vaughan, station 6021, Cat. No. 35301, U.S.N.M.), enlarged 100 diameters. The section shows the welldeveloped "coaxial" hypothallium, the smaller-celled perithallium, and the conceptacles with the coarse intersporangind tissue characteristic of the genus Lithothamnium.

PLATE 10.

Lithothamnoium isthnmi M. A. Howe.

A specimen from about one-third mile north of New Frijoles (MacDonald and Vaughan, station 6024-b, Cat. No. 35305. U.S.N.M.), natural size, showing fossil embedded in matrix, in both weathered and freshly broken surfaces.

PLATE 11.

Lithothamnium isthmi M. A. Howe.

A somewhat obliquely transverse section of a branch (specimen from about 4 miles north of Gamboa Bridge, MacDonald and Vaughan, station 6021, Cat. No. 35302, U.S.N.M.), enlarged 106 diameters.







































































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INDEX.



Page. Page.
Areha eolithothamnium ---------------4 Lithothamuniuin fosliei ---------------11
dimotum --- 5 fragilissimum ------ !
episporum-- 1-6, 12 glaciale ----------- 7
erythraeum.... 3-5 isthmi.-____ 2, 8, 10, 11, 1%S
zonatosporum 5 lichenoides ----------9, 1(0
Eulithothamnium fosliel -------------11 philippiil----------- 1
Goniolithon ------------------------3 pulclirum------------1
Lithophyllum -----------------3, 9-41,13 ramoissimum ---------1
crispatumfi--------------10 suganum ------------daedaleum -------------8p 10 vaughanlii-------2,6e-8, 13
decussatum -------------10 Mastophora melobesioldes ------------11
racemus ----------------7 Nulllpora ramosissima ---------------11
Lithoporella melobesioides -----------2, 11 Sporolithon ptychoides --------------4. 5
Litbothumnium ------------2, 8, 10, 11, 13 Stlchospora,---------------------crassum --------------7 Stromatopora compacta --------------I
curasavicum ---------1, 5






SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM Bulletin 103


CONTRIBUTIONS TO THE GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE, PANAMA, AND GEOLOGICALLY RELATED AREAS IN CENTRAL AMERICA AND THE WEST INDIES




THE FOSSIL HIGHER PLANTS FROM

THE CANAL ZONE


By EDWARD W. BERRY
Of the Johns Hopkins University, Baltimore





Extract from Bulletin 103, pages 15-44, with Plates 12-18




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WASHINGTON
GOVERNMENT PRINTING OFFICE 1918











THE FOSSIL HIGHER PLANTS FROM THE CANAL ZONE.1


By EDWARD W. BERRY,
Of the Johns Hopkins University, Baltimore.


INTRODUCTION.
It is a truism that the present floras and faunas of Central America are the result of a long series of antecedent geologic changes which might be amplified as geographic, climatic, and biologic. As the past can only be understood by means of our knowledge of the present, so, too the present can only be understood by means of our knowledge of the past. Moreover, this can never be a local problem, and this is particularly true of the Isthmus of Panama marking as it does at times the highway of communication between the terrestrial life, both animal and plant, of North and South America; at other times marking one of the paths of communication between ie marine life of the Atlantic and Pacific. Thus the history of the Central American region is of the utmost importance in any consideration of the extinct terrestrial faunas and floras of North America or 'the marine faunas that formerly flourished on the east and West coasts.
Our knowledge of the present flora of the isthmian region is based Upon Seemann's flora 2 and Heinsley's flora of Central America, supplemented by the scattered papers by numerous authors on special Copies relating to this flora. As the results of the recent Biological Survey of the Canal Zone become available, we will doubtless have a. secure basis for comparisons with antecedant floras both in this region and the area4 north and south of it.
The present distribution of plant associations is in its broader outlines governed almost entirely by the interrelations between
LR. T. Hill, who did some geological work on the Isthmus in 1895 for Alexander &gassiz, mentions lignite and fragments of fossil plants in the Culebra clays at the base t the canal cutting at Culebra station (Bull. Mus. Comp. Zool., vol. 28, No. 5, 1898), and 'he lignitic coal at Chiriqui Lagoon was studied by Dr. John Evans in 1857, who reported that the fossil plants associated with the coal were endogenous and allied to or identical with those at present growing in the vicinity." (Repts. of Expl. & Surv. for the Loca'ion of Inter-oceanic ship canals, etc., by the U. S. Naval Exped., 1875, E. P. Lull, U. S. N., commanding, Washington, 1879.)
28eemann, Flora Panamensis, Botany of the voyage of H. M. S. Herald, pp. 57-346, 852 1857.
15





16 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
topography and the prevailing winds and the resulting variations in rainfall.
The climate is now moist tropical, modified by the nearness of the two oceans, and there is therefore but slight diurnal or annual variations in temperature. So far as information is available regarding the conditions during the Tertiary, there is no evidence that can be deduced from the fossil flora or the geographical history of, j the region to indicate that the climate was very different from what it is now at any time during the Tertiary, unless we are prepared tc t assent to enormous changes in the altitude of the land, for which the data does not seem to be adequate.
The prevailing winds now come from the northeast, and as the i divide is near the Pacific Coast the major part of the Isthmus north 1 of this low divide has a heavy rainfall, as, for instance, 170 inches, 0 at Porto Bello and 129 at Colon, as compared with 90 inches at R Culebra or 71 inches at Ancon. There are two seasons-a short, relatively dry season extending from January to April and a long and relatively wet season the balance of the year with the maximum r. of precipitation from September to December. Before the clearing of the French Canal Company forests covered six-tenths of th: Isthmus, the remainder being broken forests and savannas. Evergreen tropical rain-forests of mixed angiosperms covered the entire northern watershed and part of the Darien region on the south side Some of the forests of the southern watershed are what are knowI as monsoon forests, with many deciduous species, and at high alti tudes there may be more gregarious types of forest as, for example the oak forests which are so striking a feature in the uplands o: Central America as you proceed to the northwest.
The shores are skirted with dunes abounding in Leguminosae an( Euphorbiaceae with Coco palms and Hippomane. Low shores anq tidal inlets are covered with mangrove swamps with Rhizophors," Avicennia, Conocarpus, etc. Less saline coastal marshes are cover# with Acrostichum, Crescentia, or Paritium thickets. The evergreen forest is composed chiefly of species of Sterculi~ceae, Tiliaceae, am Mimosaceae, Euphorbiaceae, Anacardiaceae, Rubiaceae, Myrtacea. and Melastomataceae, with small palms like Chamaedorea, Trithri, nax, and Bactris.
CORRELATION.
The fossil flora described in the present report is too limited fo purposes of exact correlation, which may be expected to be settle Iy the marine faunas present at most horizons in the Isthmian region Regarding the plants in the various formational units recognized i the Canal Zone by MacDonald a glance at the accompanying tab],, of distribution will show that from the oldest (Bohio) to the young





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 17

est (Gatun) plant-bearing formations there is no observable difference in floral facies, and while the plants are entirely too few for positive conclusions, and while not much variation can be expected in fossil floras of the Tropics unless after the lapse of long intervals of time or the intervention of marked changes in physical conditions, I am disposed to think that this so-called Oligocene series of formations does not represent any great interval of time.
Nearly all of the fossil plants are new, the only outside occurrences being the Hieronymia which is common to the Tertiary of Ecuador and the Palmoxylon and Taenioxylon both of which occur in the, Oligocene of the island of Antigua, and both have related types in the Oligocene (Catahoula and Vicksburg) of our Southern States Tn addition to the Hieronymia common to Ecuador there are several other elements in the Tertiary flora of the latter region that are similar to Panama forms, and it is not improbable that the coals of Loja in the Ecuadorian Andes are the same age as the so-called Oligocene series of Panama. Only one pre-Oligocene plant is recorded from Panama and the age (Eocene) rests on the stratigraphic observations of Doctor MacDonald and paleontologic determinations by C. W. Cooke. The form itself offers no intrinsic evidence of its age and might well be early Oligocene but for the fact that Doctor MacDonald collected the type stratigraphically below a bed containing a varietal form of the mollusk, Venericardia planicosta.
The chief question of interest in the correlation of these Panama beds is their equivalence in terms of the European section. Th, present flora offers no evidence on this point which must hence be determined by the accompanying marine faunas. However, in view of the traditional unscientific assumption that all of the fossiliferous beds of the Carribbean region are Oligocene in age, it is of interest to note that Douvill' 1 from a study of the foraminifera, pointed out as early as 1898, that a considerable part of the so-called Oligocene of the Isthmus was Aquitanian and Burdigalian in age; that is to say, lower Miocene according to the present conceptions of European geologists and palentologists.
In my preliminary announcement 2 of the discovery of fossil plants in the Canal Zone I stated that none of the plants recognized indicated Eocene and that they were all probably Oligocene in age. This statement was perhaps overemphasized in a desire to offset the extreme views of certain foreign paleontologists who have held that these faunas were young Miocene or even Pliocene.
The question of the exact time in the Tertiary at which connections between North and South America were replaced by marine conditions is of the utmost importance in all studies of distribution of both
IDouvilM, H. Bull. Soc. Gdol. de France, ser. 3, vol. 26, pp. 587-600, 1898.
2 Berry, E. W. Science, new ser., vol. 39, p. 357, 1914.






18 BULLETIN 103. UNITED STATES NATIONAL MUSEUM.

the marine faunas and the terrestrial faunas and floras. The floral evidence as previously stated is inconclusive. I should not, however. be inclined to consider any of the fossil plants, except one Eocene species, described in the present report as younger than Burdigalian nor older than Sannoisian (Lattorfian).


K=




6840
0 goo PQ .6845
Palmoxylon plmacites .............................. ... ... ... ...... X




6837
Ficus cuiebren is ..................................... ..................... ...... X ...... ...... .....
6840
Cuatteria culebrensis ........................... X..... .. .. ...........
Myristocophyllum pammee ......................... ..... ..... ..... .....X ............
1
6523 1 6845
Taenioxylon multiradiatum ................................ ...... I X ...... X


inga oligocaenica ............................... ..... ...... ...... . ... -.. .


68407
Cassia culebrensis ....... .......................... ...... 6840 ...... ...... X .................
Hiraea oligocaenica ................................. ...... ...... .... .. ...... ...... ...... .....
Banisteria praenuntia ................. ............. ...... ...... ............... .....
Hieronymia lehmanni ............................. ...... X .... ... X- ......... .
6840
Schmidelia bejucensis ................................. ...... ...........X
iespilodaphne culebrensis............................. ..... ..... ..X ...... ...... ....
Calyptranthes gatunen.sis ............................. X ...... ...... ...... ...... ...... ...... .....
Melastomites miconioides.............................. .. .......... X .. .....
Diospyros macdonaldi .................................................... ...x...
Rondeletia goldmani ...........................X ... ...
Rubiacites izoreoides .......................... X .. ...... ....... .

6839
Palm rays................................................................... X I mile 8: of Empfi
6837 Bridge.
Fern fragments.............................................................. 6837 cf. Acrostichum.

BOTANICAL CHARACTER.

The fossil flora at present known from the Canal Zone is extremely limited and entirely too small for either purposes of adequate cotrrelation or for deductions concerning the true botanical facies or the environmental conditions. Seventeen species are determined an two or three additional forms are tentatively recognized. Thi
paucity is especially to be regretted since it is improbable that under the existing climatic conditions as favorable opportunities for th discovery and collection of fossil plants will ever be presented a during the digging of the canal. While fossil plants were nowhere
found to be abundant in the shales, nevertheless, it is very probable that an experienced collector by working over a large amount of






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 19

material could have gotten together a much more representative collection.
The plants collected include ill-defined fragments of one fern, two undertermined species of palm, represented by fragments of foliage. and a third represented by petrified stems, and 16 dicotyledons, of which two are represented by fruits and the balance by leaves.
Among the Dicotyledonae there are representatives of the orders Urticales, Ranales, Rosales, Geraniales, Sapindales, Thymeleales, Myrtales, Ebenales, and Rubiales. Orders conspicuous in the existing flora of the Isthmian region unrepresented among the fossils are the Arales, Poales, Cyperales, and Orchidales among the Monocotyledonae, and the Campanulales and Personales among the Dicotyledonae.
The following 14 families are represented by fossils in Panama: Moraceae, Anonaceae, Myristicaceae, Mimosaceae, Caesalpiniaceae, Papilionaceae, Malpighiaceae, Euphorbiaceae, Sapindaceae, Lauraceae, Myrtaceae, Melastomataceae, Ebenaceae, and Rubiaceae. Only the last, with two species, is represented by more than a single species. When so sparse and evenly distributed a representation of the families is present in a fossil flora, it is an indication that after allowing for some accidents of preservation, those families represented may be regarded as the-most abundantly represented in the Tertiary flora of the region, and in this respect there is a very great similarity to the existing flora of the Isthmian region. The present forests of Panama are made up principally of species of Arecaceae, Moraceae, Mimosaceae, Papilionaceae, Sterculiaceae, Tiliaceae, Euphorbiaceae, Anacardiaceae, Myrtaceae, Melastomataceae, and Rubiaceae. The only ones of this list not found fossil are the Sterculiaceae, Tiliaceae, and Anacardiaceae, and as these three families are all abundant in the much more complete floras from the Tertiary of the southeastern United States, it is safe to assume that they were also present in the Tertiary flora of Panama. The mainly herbaceous families abundant in the Recent flora, which are hardly to be expected in the fossil flora, are the Poaceae, Cyperaceae, Orchidaceae, Araceae, and Compositae.
The bowers of wild figs of the existing flora are represented by a small-leafed species of Ficus from two localities in the Culebra formation. The family Anonaceae, which has numerous species of Anona and Guatteria in the Recent flora of Central America, is represented by a fine large species of the latter genus which is not uncommon in the Gatun, Caimito, and Culebra formations. Guatteria contains about 50 existing species of tropical shrubs and trees of varying habitats and exclusively American, and has not been previously recognized with certainty in fossil floras. Anona is abundant






20 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
I
in the Eocene and Oligocene of our Southern States, but Guatteria has not been recognized.
The Myristicaceae is represented by an infrequent species of f Myristicophyllum in the Culebra formation, and in this connection it ( is of interest to note the presence of fruits and seeds of Myristica in s the uppermost Eocene of Texas suggestive of the subgenera Virola c and Compsoneura. both of which occur in the Recent flora of Central America. The Leguminosae have three fossil species. The Mimosa- e ceae, which are very abundant in the existing forests of Panama, are I represented by a fossil species of Inga, a large genus of tropical trees S with upward of two-score species in Central America, nearly half I of which are recorded from Panama. Inga is well represented in 2 the abundant Eocene floras of our Southern States, and it is of in- I terest to note the resemblance between the fossil species from Panama and a species described by Engelhardt from an unknown Tertiary horizon in Ecuador.
The Caesalpiniaceae is represented by a single species of Cassia, e large genus not only in the Recent equatorial floras but well represented in most fossil floras from the Upper Cretaceous to the present
The Papilionaceae, very abundant in the existing flora of Panama I is supposed to be represented by the petrified wood of a large tre~ referred to the genus Taenioxylon and found in the Cucuracha Culebra, and Bohio formations.
The family of Malpighiaceae is represented by the genera Hiraec and Banisteria. The former has about 30 recent species, exclusively American, ranging from Mexico and the Antilles to tropical Brazi and Peru, and it is represented by a fossil species in the Eocene o:I the Mississippi embayment. Banisteria contains about 80 existing species, mostly climbing shrubs. It is at present confined to the American tropics, but appears to have been present in Europe as wel as in the southern United States during the Tertiary.
The Euphorbiaceae, abundantly represented in the present forest of P:unama, is represented in the Caimito formation by a species o: Hieronyma apparently identical with one described by Engelhard from the Tertiary of Ecuador. Hieronymiia, not otherwise knowli in the fossil state, contains about a dozen existing species which ar confined( to tropical America, where they range from Mexico and tht West Indies to Brazil.
The Sapindaceae, abunmdant in all fossil floras from the Uppe (Cretaceous onward, and exceedingly abundant in the Tertiary flora of the Mississippi embayminent, is represented in the fossil flora oPanama by a species of Schmidelia found in the Caimito and Culebr format ions. Schmidelia has a large number of existing species ii the equatorial regions of both hemispheres and, except for petrifie





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 21

material from the island of Antigua, it has not previously been recognized in the fossil state.
The family Lauraceae, so extensively represented in the Tertiary floras of the Mississippi embayment and in the Recent tropical flora of South America, is represented at Panama by a single fragmentary species which is referred to Mespilodaphne. The latter has numerous moder species in the tropics of America and Africa. The Myrtaceae, one of the abundant families in the existing forests of tropical Anerica, has a fossil species of Calyptranthes at SPanama. This genus has about 70 exclusively American existing species ranging from Mexico and the West Indies to southern Brazil.
Hemsley records 7 recent species from Central America, of which 2 are found on the Isthmus. It is also represented in the lower Eocene of the Mississippi embayment. The abundant, both Recent i and fossil, representatives of the allied genera Eugenia and Myria
7 have not been recognized in the fossil flora of the Isthmus.
The Melastomataceae, an immense tropical family in the existing
flora and very abundant throughout Central America, has a single
fossil species in the Culebra formation.
iThe family Ebenaceae, usually abundant in fossil floras from the Upper Cretaceous onward, and with a large number of species in tropical America, is represented on the Isthmus by the petrified fruits of a species of ebony (Dio pyros) known to be from an older
horizon (Eocene) than the.balance of the known fossil flora.
The Rubiaceae, a prominent family in the existing flora of Central
America, where according to Wallace (1911) it ranks fourth in size with 146 species, is represented by two fossil species, both found in 1 the Gatun formation. These are referred to Rondeletia and Rubiacite8.
The former has not heretofore b~en found fossil. It includes about 70 existing species of a variety of habitats, confined to the American tropics and chiefly massed in the Antilles and Central America. Rubi'Wltes is represented by a fruit which is apparently referable to the tribe Ixoreae, now confined to the tropics of both
-hemispheres.
TERTIARY ECOLOGY.
The restricted variety and fragmentary condition of the fossil
plants thus far collected inhibits a detailed discussion of the probable ecology of the Tertiary flora. In so far as climatic conditions re concerned the Tertiary plants indicate an abundant rainfall and relatively high equable temperatures such as prevail at the present time in the Hill country and Coastal Plain of the Istu s. There is no indication of upland vegetation. None of the fossil plants indicate





22 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

mountains sufficiently high to harbor that mixture of temperate types 0' such as is seen at the present time in the mountains of Central America, as, for example, above 6,000 feet in Costa Rica. There was plenty of opportunity for the introduction of such types had the climate been propitious, so that I would infer that the Tertiary relief was slight, that is under 5,000 feet and probably much less than this, although there is no evidence to warrant precision of statement.
On the other hand, the collected floras do not furnish any traces of the characteristic vegetations of low muddy shores, although types like Rhizophora, Avicennia, Conocarpu,, Laguncularia, etc., were already in existence in Eocene times as we know from their presence in the Mississippi embayment of that time, where they were undoubtedly derived from the south. I do not infer that these costal types M were absent in the Tertiary flora of the Isthmus. On the contrary they must have been present; but no traces of them have been discovered except the traces of Acrostic.hum in the Culebra formation.
The bulk of the fossil plants clearly belong to the evergreen rain
forests and they have the appearance of having been washed into !1 the basins of sedimentation by streams. None of the lithologic specimens that I have seen from the Isthmus indicate autochonous swamp deposits either of coastal or valley situations and I picture the flora 3 as one of a humid tropical character covering a country of low hills. This is of necessity a tentative conclusion and perhaps even such general deductions are unwarranted because of the very limited data with which I have had to deal.

FLORA OF THE CANAL ZONE.
Arecales:
ArecaceaePalnwmxylon palmacites (Sprengel) Stenzel. Urticales:
MoraceaeFicus culebrenwis, new species.
Ranales:
AnonaceaeGuatteria culebrensis, new species.
MyristicaceaeMyristicophyllum panamense, new species. Rosales:
LeguminosaeTaenioxylon multiradiatwm Felix.
Inga oligocaenica, new species.
Cas8ia ebrensi8, new species.


Il





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 23

Geraniales:
Malpighiaceae-Hirea oligocaenica, new species.
Banisteria praenuntia, new species.
EuphorbiaceaeSHieronymia lehmanni Engelhardt ? ISapindales:
Sapindaceae-,Schmidelia bejucensie, new species. Thymeleales:
Lauraceae-Mespilodaphne cdulebremsis, new species. Myrtales:
SMyrtaceae-Calyptranthes gatunensis, new species.
MelastomataceaeMelastomdtes miconioides, new species. Ebenales:
Ebenaceae- Diospyros macdonaldi, new species. Rubiales:
Rubiaceae-Rondeletia goldmani, new species.
Rubiacites ixoreoides, new species. Fern fragments of Acrostichum. Pal rays.

SYSTEMATIC PALEOBOTANY.

PTERIDOPHYTA.

Order FILICALES.

FERN FRAGMENTS OF ACROSTICHUM.

The material from the Culebra formation, one-fourth mile south of Empire Bridge, contains several obscure fragments of large simple fern pinnules with reticulate venation strongly suggestive of Acrostichum, but too incomplete for identification. The genus now principally represented by the cosmopolitan tropical tidal marsh species Acrostichum aureim is abundant in the Eocene and Oligocene of Sboth America and Europe, and is especially characteristic in the Jackson, Catahoula, and Vicksburg of our Gulf States.





24 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

SPERMATOPHYTA.

Order ARECALES.

Family ARECACEAE.
PALM RAYS.

The broken rays of apparently two species of palms occur sparingly in the Culebra formation at the locality one-fourth mile south of Empire Bridge. These are too incomplete for even tentative generic determination.

Genus PALMOXYLON Schenk.

Group LUNARIA.

PALMOXYLON PALMACITES (Sprengel) Stenzel.

Plate 12, fig. 1.
Endogenites palrmacites SPRENGEL, Commentatio, p. 39, figs. 6, Oa, 1828.
Fasciculites palmacites COTTA, Dendrol., pp. 49, 89, pl. 9, figs. 1, 2, 1832.UNGER in Martius, p. 59, tab. geol. 3, fig. 6, 1845.
Palmacites dubius CORDA, Beitrage, p. 42, pl. 22, 1845.-SCHIMPER, Pal
Vgft., vol. 2, p. 513, 1870; Handbuch, Abst. 2, p. 887, 1892.
Palmacylon tenerum FELIX, Foss. Hdlzer Westindiens, p. 26, pl. 4, fig. 1
1883.-SCHENK in Zittel.
Palmoxylon palamacites STENGEL, FOSS. Palmenhilzer, p. 245, pl. 20, fig. 252
1904. i
Description.-Fibro-vascular bundles small, very numerous, closel; spaced, orbicular or ovate in cross section, uniformily distributed a a rule, 0.60 mm. to 0.75 mm. in diameter, and rarely, if ever, tha distance from one another. Auxiliary bundles absent.
Sclerenchyma portion excavated more or less deeply to receive th vascular portion, which is often nearly equal to it in size. Occa sionally a thin zone of sclerenchyma entirely surrounds the vascular portion. Sclerenchyma fibres small, isddiametric, greatly thickened of nearly uniform size, about 0.035 mm. in diameter. Vessels vanri able in size, ranging from 0.072 mm. to 0.18 mm. in diameter, usuall two large vessels and either none or several small vessels on the sid : way from the bast in each bundle. The phloem portion in genere destroyed and represented by a disorganized cavity between the ves sels and the bast.
The ground mass of the stem consists of thin walled parenchym without intercellular spaces. The cells are small, isodiametri rounded pentagonal or hexagonal except where there are but one two rows between closely adjacent bundles, in which case they ai






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 25

narrowly compressed and elongated parallel to the sides of the bundles. Their diameter varies from 0.035 mm. to 0.10 mm. Scattered through the stem parenchyma are darker cells which in polarized light appear to be gum cells. They are slightly larger than the parenchyma cells, being from 0.072 mm. to 0.108 mm. in diameter.
Occasional bundles are seen to be branching. These are the fasciciuli flbroductores or Kreuzungsbiindel. it This species was first recognized by Sprengel in 1828, who referred it to Endogenites; Cotta four years later transferred it to Fasciculites, and Corda in 1845 referred it to Palmnacites. When Felix came to publish on the Antigua woods in 1883 he recognized this species, but in describing it under the genus Palnwxylon which had been proposed by Schenk only a year or two before he took the liberty of giving it the new name of tenerum, which under the rules of nomenclature has no standing as Stenzel recognized in print in 1904.
The specimen from Panama is small and may be from near the periphery of a stem, although in the group Lunaria there is little diference between the central and peripheral regions. In the size, outline, and crowding of the fibrovascular bundles as well as in the character of the parenchyma of the groundmass the present species greatly resembles Palmoxylon integrum described by Felix 1 from Cuba and considered by StenzelJ as merely a variety of the Antiguan species Palmnwylon antiguege (Unger) Felix.3 It differs from that species in altogether lacking the numerous auxiliary sclerenchyma bundles which are so well marked in Palmoxylon integrum. A further difference is the presence of gum or mucilage cells which are fairly numerous in the Panama specimen of Palmoxylon palmacites and which might upon a merely superficial examination be mistaken for auxiliary sclerenchyma bundles. Among the Oligocene species of Palmoxylon from the southern United States Palmoxylon missisSippen~e Stenzel is very similar to the present species.
Other described fossil species which show more or less resemblances are Palmoxylon stellatum, aschersoni, variabile, and ceylanicum. The nearest affinity among recent palms is not determinable in the present state of our knowledge of the anatomy of the latter. The present type of structure is commonly known as the Cocoslike type.
For some unknown reason the upper Eocene and lower Oligocene in southeastern North America abounds in silicified palm wood. Palm leaves are often very abundant in the Wilcox and Claiborne Eocene and in the Apalachicola Oligocene; but all of the petrified
'Felix, Foss. H6lz. Westindiens, p. 24, pl. 5, fig. 2, 1883.
2 Stenzel, Foss. Palmenh6lzer, p. 154, pl. 1, figs. 1-10, 1904.
8 Felix, Foss. H6lz. Westindieus, IV. '22 pl. 4 ftig, 15
SStenzel, Foss. Palmenhileir, p. 241, pl. 21, 4gsi 254 -26.





26 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

palm wood in our Gulf region is confined to the Jackson or Vicksburg groups. I.et
The island of Antigua, celebrated for at least a century for its.0 petrified woods, has furnished at least seven species of petrified ai palms, five of which were known to Unger as early as 1850, and on( by was figured by Witham in 1833. These also are of Oligocene age. There are two additional Oligocene species described from the West Indies without definite information as to exact locality, and there is also a species from Trinidad and another from Cuba. The Oligo0cene species at present known from the southern United States are seven in number, four of which have not been found outside of that region, while one or possibly two are common to Antigua, and a third has been reported by Felix from Southern Mexico.
Occurrence.-Cucuracha formation, green clays, Gaillard Cut (loc. 6586). Collected by D. F. MacDonald.
Collection.-U. S. National Museum. Cat. No. 35310.

Order URTICALES.
Family MORACEAE.
Genus FICUS Linnaeus.
FICUS CULEBRENSIS, new speeles.
Plate 13, fig. 1.

Description -Leaves of relatively small size, broadly oblonglanceolate in general outline, apex acute but not extended or cuspidate. Base bluntly pointed. Margins evenly rounded. Texture-. coriaceous. Length about 8 cm. Maximum width, in the middle part of the leaf, about 2.15 cm. Petiole short, stout, and curved. Midrib stout and prominent on the under surface of the leaf. Seeondaries thin, very numerous, evenly spaced, subparallel; they diierge from the midrib at wide angles averaging about 75 degrees, pursue an almost straight outward course, their ends being conitocted well within the margins by regular flat arches formed by their abrupt camptodrome endings. Tertiaries obsolete.
This is an especially well-marked species of the lanceolate leafed section of Fic-'u, and it may be matched by a number of still existing species found in the American tropics. Among such a large number of both existing and fossil forms detailed comparisons are not especially pertinent. Two comparisons that seem significant are the resemblance of the present form to Ficus new tonensis Berry of the I ~ppCr (laiborne of the Mississippi embayment and to the forms from, the Sannoisiar of Paor'ing in the Tyrol which Ettingshausen1 refers
,fltttngShausen, Tert. Fl. von 1lacring. p. 41, pl. 10, figs. 6, 8, 1858.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 27
3- to Ficus jync Unger, but which appear to me to be decidedly different from Unger's type.
Occurrence.-Culebra formation, upper part. East wall of the e Gaillard cut just north of Canal Commission station 1760 (collected w by M. I. Goldman).


Order RANALES.
Family ANONACEAE.
it Genus GUATTERIA Ruiz and Pavon.

r GUATTERIA CULEBRENSIS. new species.

Plate 13, fig. 2.
Description.-Leaves of large size, broadly ovate in general outline, with a narrowed slightly decurrent base and a narrowed and extended acuminate tip. Length about 20 cm. Maximum width, approximately midway between the apex and the base, between 6 cm. and 7 cm. Margins entire. Texture coriaceous. Petiole short and stout, enlarged proximad, about 2.25 cm. in length. Midrib stout and promient. Secondaries mediumly stout and prominent, about ten opposite to alternate pairs diverge from the midrib at angles ranging from 450 to 60, sweeping upward in regular ascending subparallel curves, camptodrome in the marginal region. Tertiaries,
1 Where visible, percurrent.
The present is one of the more abundant and better preserved i forms from the Canal Zone, but the large size of the leaves usually l results in fragmentary specimens, the tip being almost invariably missing. The species shows great similarity with various existing forms of Anonaceae. It is very close to Anona maregravii Martius of Venezuela, French and Dutch Guiana, and Brazil (Bahia and Pernambuco). It is, however, among the various species of Guatteria
that the closest homologies are found. The latter genus contains about fifty species of shrubs and trees, exclusively American I and found in Mexico, Central America, tropical South America, and in the northern Andes. The fossil may be compared with a large number of the existing species, as for example Guatteria ouregon Dunal, a large tree of the Carribbean islands and equatorial South America. Guatteria dolichopoda De Candolle or G. grandiflora De Candolle iof Central America.
The family Anonaceae contains about 700 existing species, distributed among about 48 genera, only two of which are present in North America. The family is practically confined to the Tropics,
'The Asiatic species of various authors are referred to the genus Polyalthia.
83700-18g-Bull. 103-3




*!

28 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

a single Australian species, and the North American genus Asimin, with 6 or 7 species being the only conspicuously extratropical forms. The area of maximum representation is southeastern Asia and the adjoining region of Malaysia, for while only 16 genera are confined to this region it contains over 350 species, and six additional genera,
(Miliusa, Uvaria, Polyalthia, Oxymitra, Melodorum, and Poporvia), with a total of over 250 species have the bulk of their species in this area. Only a single genus is confined to Australia, and the bulk of the Australian species are to be regarded as migrants from the preceding area. There are upwards of 100 species and 6 peculiar genera 01 in tropical Africa; and America has about 200 species and 10 peculiar genera. These are all confined to the Tropics, except for a species of Anona, which reaches the coast of peninsular Florida, and for the genus Asimina, with six or seven species of shrubs and small trees of the south Atlantic and Gulf States. One of these, .tsimina triloba Dunal, is hardy as far north as New York, and has the distinction of growing the farthest distance from the Equator of any existing member of the family. The fossil record of the Anonaceae is very incomplete, only the genera Anona Linneaus and Asimina Adanson,. being known with certainty. Both of these genera are present in the flora of the Wilcox group of the Mississippi embayment.
The genus Guatteria has not, so far as I know, been heretofore found fossil, except for a doubtful species described by Hollick from the Upper Gretaceous of Marthas Vineyard and Long Island. The genus Uvaria Linnaeus has a Pliocene and three Pleistocene species on the Island of Java, and the genera Melodorum Dunal and Mitrephora Blume are both represented in the Pleistocene of that island.
The genus Anona has from fifteen to twenty fossil species, five of which are also represented by seeds. The oldest is a species described from the Dakota sandstone. There is a second species in the late Cretaceous or Early Eocene of the Rocky Mountain province. TheI flora of the Wilcox affords a glimpse into the true stage of evolution of Tertiary floras in that expanded belt of the American equatorial region which was the center of radiation of so many recent types. There were three exceedingly well-marked species of Anona along the Wilcox coast and their leaves are very common at some localities, although no seeds have as yet been discovered. I assume that these Wilcox forms had habits similar to those of the majority of the exi.ti1g species, exemplified by our Florida Anova glabra Linnaeus, or pond apple, which frequents shallow fresh-water swamps, low shady hammocks, or stream borders near the coast. Other species occur in the low coppice association or on edges of brackish swamps (m the Bahmas. fhe cultivated species, as, for example, the Ameri ionn 1tio retwuata Linnaeus. which is planted in Guam, often


Ii





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZON*. 29

spreads naturally along the inner beaches, while attempts to introduce others of the most highly esteemed American species in the Orient have failed. From its prevalence among the existing species the habit of growing in wet, shaded soils is evidently an old one, and since the Wilcox Anona8 are associated with a strand flora the assumption that they grew on the inner beaches or the shaded and more swampy edges of lagoons possesses every degree of probability.
In the pipe clays of Alum Bay which were contemporaneous with the Wilcox there are two species of Anona, and Engelhardt has described two species from the Eocene or Oligocene of Chili. The Oligocene record shows a species in France and a second in Saxony. In the Miocene there are two species each in England, Styria, and Croatia, and one each in Bohemia, Colorado, and Transylvania. There is one each in the Pliocene of France and Italy, showing how modern was their extinction in the south of Europe.
The genus A-simlna has only four or five recorded fossil species. These are all American except for a form from the Pliocene of Italy which has been referred to this genus, although I suspect that it presents Anoaa., since Asimna appears to have originated and been confined to the Western Hemisphere. The oldest known species is based on foliage which is found in the basal Eocene of the Rocky IMountains (Denver formation) and of the embayment (Midway Group). There is a single species based on a seed from the basal Wilcox and no other records except a form close to the modern from the late Miocene of New Jersey (Bridgeton sandstone) and the occurrence of the existing Asimnina' triloba Dunal in the interglacial beds of the Don valley in Ontario. There are 17 existing species of Anona recorded from Central America, six of which are known from Panama. Hemsley records 11 species of Guatteria from Central
America, at least two of which occur in Panama.
Occurrence.-Culebra formation, upper part. East wall of Gaillard Cut just north of Canal Commission station 1760 (collected by M. I. Goldman). Gatun formation. Gatun borrow pits (collected by M. I. Goldman). 7 miles northeast of Bejuca near Chame t=Caimito formation) (collected by MacDonald).
Family MYRISTICACEAE.
Genus MYRISTICOPHYLLUM Geyler.
MYRISTICOPHYLLUM PANAMENSE, new species.
Plate 13, fig. 3.
Ieseviption.-Leaves ovate or ovate lanceolate in outline with pointed apex and base, entire, evenly rounded margins, subcoriaceous in texture. Length about 9 cm. Maximum width, midway between the apex and the base, about 3.3 cm. Petiole slender, about 8 mm. long. Midrib slender. Secondaries thin, about 8 subopposite ascend-






30 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

ilig sutbparallel pa-ir-s thiey diverge from the miidrib at acute angles I aiili are st~lbparallel with the lower lateral margins, eventually camip- z todrome. Tertiaries obsolete.
Tlii~s species is unfortunately represented by fragmentary remains Iiadeqluate for conclusive identification. The genus, Jlyristica Lin- f Idells, conltainis about two score existing species, rather more thanl half being A4kiericall tropical forms, now often segregated into several genera. Manyv are insular and coastal forms, Schimpet' recording 4 a sI-)e(1es in thie l~ndonialayan strand flora and several species ranging t, eastward in the Pacific to the Fiji, Tonga, and Samoan Islands, and 1i their fruits are recorded by both Gaudichand and Guppy in the seat drift, although the oriental species are normally'distributed by fruit pf)IgQous (Mloslev. Heislev. Gu1ppy). t
D~e Caiidolle and"Mwiel 1)oth considered, the, foliage, especially the '
VC111,1tio)t offeilO( the best criteria, for differentiationbuinteii
-IbZ7e(Iie of comiparat ive material, and the incomplete character of theJ IPanaina, fossil it is not possible to apply these criteria. The Am-erican Recent species number about '25, and these are mainly South American in their distribution, although the sections or genera. Vh'iolatJ Aublet and Cornpsoneura De Candolle both occur in Central America.w
Thie distribution of the Recent species in tropical America. Asia, and. Africa is conclusive evidence of a Tertiary history, although this ff evidence, is pr-actically unknown. Gieyer' described two forms of leaA f ragnients fromn the Miocene of Labman (Borneo) and Engelhardt 2) third from the Tertiary of Ecuador and Chile. The most concluisivc, i evidence of their Tertiary radiation is furnished by the character-isti(, 0, fruits described recently by the writers3 and preserved in the windi-4 blown stands of the uppermost Eocene of Texas.
Oe.-cv)rrnce.-Culebra formation (upper part). East wall of Gail I lard (Cit just north of Canal Commission station 1760 (collected bil ~J At 1. Goldman).
Order ROSALES.
Superfamily LEGUMINOSAE.
Genus TAENTOXYLON Felix.
TAENIOXYLON MULTIRADIATUM Felix.

TI(/I(-) UJ.Vylfill ii: I? Itir(I di HI it I) i V1~ XA, 1 )ie f oss il I e 1 ilzer 11Wes'tirdien"
S:1i11i11I. IJ;iewoi1. Abl.. sec. 1, Hl'i 1, p. 11, Id. 1. fg.10, 11; p1.
figl. 1W. 1M.1S
jiv~floKei-s(, sW(tl.-Iii a radial dlistaince of 5i ciii. there ar-e n1,
(leliilie annu111al or seasonal1 rings. In cer-tainl zones thle vessels ar Ir
'4 ~4y 1 1. JI T., I uyal Expodit ion, vol. A, p). 498, pl. 3,figs. %-G. 1887.
V n~1I111id, H., Ablh. Selitck. Naturf. (ese11-ch., vol. 16, 1p. 66;, lit. G, fig.- 9;p1.
~ W-ry.ICW.. Allir. .Io irn so, tr. 4, vol. 42, pp. 2,41-2-47i, ligs, 1-6. 1910.





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 31

S larger, more generally compound, and closer together, and in other z ones they are more distant, slightly smaller, and prevailingly single. NIo changes are observable in the other elements and there is no regular alteration of vessel rich and vessel poor areas nor any change from so-called summer to spring wood such as characterizes the trees I of the temperate zone.
di Vessels single or two, three, or four together in radial rows (an 4 anomalous group of five vessels in juxtaposition is shown in the deg tailed drawing). Outline of single vessels elliptical, those in groups d flattened on one or both sides by mutual compression; their tangena tial diameter ranging from 0.10 mm. to 0.14 mm.; their radial diami eter ranging from 0.12 mm. to 0.16 mm.. exceptionally large ones up to 0.22 nun.; their n alls thick. 0.0067 mm. to 0.01 mm. in thickness, e clearly showing the numerous small pits in section. Vessels free quently filled with gum. Vessels usually surrounded by one to three
e layers of rounded or more or les compressed thin-walled wood parenn chyma, somewhat variable in amount in different parts of the stem I- and tending to form tangential bands. Prosenchyma very abundant,
a the elements polygonal, small, somewhat smaller than those of the i. wood parenchyma, and thick walled. Rays very numerous, one or two cells wide as seen in transverse sections, flexuous in their courses since they are bowed out around the large vessels and approach more for less in the radial intervals between vessels; from 0.10 mm. to a 0.20 in. apart, averaging nearer the former than the latter figure. e The ray cells toward the ends of the rays which appear to be those usually seen in the several sections examined are not elongated radi- ally but are nearly isodiametric and about 0.02 mn. in diameter.
Radial setio.-The radial section shows the close set, fine, trans1- versely elongated pits of the vessels which have simple perforations. SThe wood parenchyma is septate, the cells being about 3{ times as long as wide with large.simple pits. The rays are of variable height. from 9 to 17 cells. They are seen in radial view to consist of a central series of radially elongated cells with numerous fine simple pits, above and below which is a series of longitudinally elongated cells, beyond which are one or two rows of isodiametric cells which are regularly hexagonal in this view.
Tangential section.-The tangential section shows the uniform close set fine pitting on all the walls of the vessels, the relative short length and the large simple pits of the adjoining septate wood parenchyma. The rays are seen to be very numerous, and separated by but few rows of flexuous prosenchyma; they are lenticular in outline and of variable height, one or two rays of terminal cells (those which are hexagonal in outline in the radial view) are single; then come one to three biseriate rows (those longitudinally elongated in the






32 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
radial view); toward the median region the rays are three or four cells broad (the radially elongated cells in the radial view).
Felix states that in the Antigua material the rays were usually biseriate, while uniseriate and triseriate rays were rare. I do not know the extent of his material, but in the case of that from Panama I had but few radial sections cut. Ray cells frequently filled to a greater or less degree with gum.
Remarks.-Fragments of the wood of this species are very common in the collections from Panama, but a good deal was rather badly decayed before petrification. That which has formed the chief basis for the foregoing description and all of the photographs and drawings is beautifully preserved. The species is clearly identical witl: 'd the type, as very insufficiently described and illustrated by Felix One highly ferruginized and fairly well preserved quadrant of a trunk indicates a large tree, with a diameter of at least 25 cm.
The genus Taenioxylon was established by Felix in 1882 witl T. varians from Antigua as the type. He has since described 7 addi tional species including 2 additional from Antigua, 1 from southern 11 Brazil, 1 from East Indies, 1 from Philippines, 1 from Caucasus and 1 from the Swabian Alps. All are of Tertiary age and shove resemblances to various members of the 3 Leguminous families Caesalpiniaceae, Miomosaceae, and Papilionaceae. Felix consider ( the present species to be a member of the Papilionaceae, and it agree 1 entirely with Solereders account of the anatomy of this family. Th k two kinds of ray cells described have, according to Saupe, beeshown to occur in the following tribes in this family, namely th E Podalyrieae, Genisteae, Galegeae, Hedysareae, and Sophoreae. Wit1 out much recent comparative material, which is unavailable, it impossible to allocate the present species more definitely within th extensive family.
Occurrences.-Bohio formation, middle Bohio Ridge (poorly pro served) quadrant of a large trunk indicating a tree with trunk I least 25 cm. in diameter. Cucuracha formation, upper part. Gree clays of Gaillard Cut (locality 6845) Oligocene limestone. Orb toidal limestone, 2 miles north of David (locality 6523) (all aboN collected by D. F. MacDonald). Culebra formation, upper pai Near top of big slide, just north of Culebra. Collected by M. Goldman (figured material).
Collections.-U. S. National Museum, Johns Hopkins.Universit Family MIMOSACEAE.
Genus INGA Willdenow.
INGA OLIGOCAENICA, new species.
Plate 16, fig. 2.
Description.-Leaflets rather above medium size, elliptical-ova and very inequilateral in general outline. Apex abruptly acute, n






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 33

Tended. Base very inequilateral, truncate or ascending on one ide and wide and cordate on the other. Margins entire, full. TexY ure subcoriaceous. Length about 8 cm. or 9 cm. Maximum width, t or slightly above the middle, about 4 cm. Petiolule curved, short 'd stout, about 3 mm. long. Midrib stout, greatly curved. Secundaries thin, five or six pairs, angles of divergence and courses arious, all ultimately camptodrome; lower pair opposite, from the op of petiolule; they diverge from the midrib at angles of about 45 egrees, curving slightly outward and then ascending, parallel with e respective margins; the one in the narrow side of the lamina rches along the margin in a brochiodrome manner; the one in the ide side of the lamina sends off on the outside a series of regularly aced camptodrome tertiaries. Tertiary venation for the most art obsolete.
This characteristic species may be compared with Inga densiflora ntham,' Inga edulis Martius.2 Inga marginata Willdenow, or Inga pdeciosa Spruce' and with various other of the larger-leafed species f Inga in the American Tropics to which region the 212 of its ting species of shrubs and trees are confined. It may also be
0mpared with a number of tropical American species of Cassia, as, or example, Casia ruseifolia Jacquin. About fiften fossil species have been referred to Inga. These elude three from the Upper Cretaceous, two European, and one I orth American. There are also two or three species in the Oligoene of Europe, one in the Pliocene of Bolivia, two in the Tertiary Af Ecuador, and one in the Tertiary of Colombia, four well-marked I pecies in the Lower Eocene of the Mississippi embayment (Wilcox
roup) and one in the middle Eocene of that region (Claiborne hroup). The Panama species is not especially close to any of the oregoing. It is nearest, however, to Inga latifolia, described by agelhardt 6 from the Tertiary of Ecuador, differing in its broader Aorm and more inequilateral base.
Pittier records 14 existing species of Inga, from Panama." Hemsy lists 35 species in his flora of Central America, or which number .8 are recorded from Panama.
Occurrene.-Lower part of Culebra beds one-fourth mile south of IL mpire Bridge. (Collected by D. F. MacDonald.) U.S.G.S. 6837.
Type.-Cat. No. 35311, U.S.N.M.
SBentham, Trans. Linn. Soc. Lond., vol. 30, p. 617, 1875 (Peru).
'Martius, Flora, vol. 20, Beibl., p. 113. 1837 (Brazil).
Willdenow, Sp. Pl., vol. 4, p. 1015, 1806 (Venezuela).
Spruce, in Bentham, Trans. Linn. Soc. Lond.. vol. 30, p. 620 (Brazil).
Engelhardt, H., Abh. Senck. Naturfor. Gesell., vol. 19, 1895, p. 20, pl. 2, figs. 11, 12.
Pittler, H., Cont. U. S. Natl. Herb.. vol. 18, pt. J. pp. 218-223, 1916.






34 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

Family CAESALPINIACEAE.

Genus CASSIA Linnaeus.
CASSIA CULEBRENSIS, new species.

Plate 16, fig. 1.
*
Description.-Leaves obviously pinnately compound. Leaflets ovate, slightly inequilateral and falcate, with an obliquely acuminate, practically equilateral tip, and an acuminate markedly inequilateral base. Length about 6.25 cm. Maximum width, about midway between the apex and the base, 2.75 cm.; one side of the lamin 15 mm. wide, the other 12.5 mm. wide. Texture mediumly coriace ous. Petiolule reduced to a thickened proximal part of the midril extending but 1 mm. below the point of junction of one margin an( about 2.5 mm. below the point of junction of the opposite margin Margins entire, evenly rounded and full. Midrib relatively thin, no prominent, curved. Secondaries thin, numerous, about 10 suboppo site to alternate pairs; they diverge from the midrib at wide angles about 700 in the middle part of the leaflet, are nearly straight regu larly spaced and subparallel in their outward course for two-third of the distance to the margin where the principal ones fork to joi: in rounded arches the similar branches of adjacent secondaries; th secondaries in the apical and basal portions of the leaflet are regu larly camptodrome; those toward the tip of the leaflet more closel ( spaced. Marginal tertiaries camptodrome, internal tertiaries mostl obsolete.
This type in its general form and the character of its base an
petiolule indicates that it is a leaflet of a pinnate leguminous lea I Its general appearance suggests comparisons with the gene Sweetia, Myrocarpws, Toluifera, Cassia, and Sophora-the first thr( confined to tropical South America and the last two cosmopolitan i the existing flora. While the evidence is not conclusive, I prefer t 6 consider it more closely allied to Cassia than to the other gene I mentioned, particularly as the venation characters are such as I has T considered referable to Cassia in my studies of the fossil floras of t southern United States. No species related to the Panama form known from the Oligocene of the United States.
The modern species of Cassia are very numerous, upwards of 4 having been described. They comprise herbs, shrubs, and trees < varied habitats in the warmer parts of both hemispheres, particular tropical America. The fossil species are also numerous and tl generic history goes back to near the base of the Upper Cretaceovu The genus has been continuously represented in the warmer parts






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 35

America from the time of deposition of the Tuscaloosa sediments of Alabama to the present.
Occurrence.-Culebra formation, lower part, one-fourth mile south f Empire Bridge (collected by D. F. MacDonald) U.S.G.S. 6837:
Type.-Cat. No. 35312, U.S.N.M.

Order GERANIALES.
Family MALPIGHIACEAE.
.Genus HIRAEA Jacquin.
HIRAEA OLIGOCAENICA, new species.
4Plate 17, fig. 1.
Description.-Leaves relatively large, ovate-lanceolate in outline, alcate, with an equally cuneatly pointed apex and base. Margins ntire, evenly curved. Texture subcoriaceous. Length about 9.5 m. Maximum width, at or somewhat below the middle, about 3.5 m. Petiole short, stout, about 3 mm. in length. Midrib stout, flexous. Secondaries thin, regularly spaced, about 9 pairs, prevailingly Iternate; they diverge from the midrib at angles of about 450 and eep upward in regular subparallel slight curves, and are camptoaome in the marginal region. Tertiaries obsolete. This genus, which has well characterized leaves, has seldom been r cognized in the fossil state. One species' is not uncommon in the lower Eocene of the Mississippi embayment, and Ettingshausen2 has recorded, but not described, a second species from the Ypresian !I Alum Bay, England.
The existing species number between 25 and 30 and are exclusively American, ranging from Mexico and the Antilles throughout Central
d northern South America to the Peruvian tropics.
The present fossil species is not unlike Hiraea wilcoxiana Berry3 om the lower Eocene of Tennessee and is closely comparable with e existing Hiraea chrysophylla Jussieu of the northern coastal reon of South America.
Occurrence.-Caimito formation 7 miles northeast of Bejuca (U.S.G.S. station 6840). Collected by D. F. MacDonald. Type.-Cat. No. 35313, U.S.N.M.
Genus BANISTERIA Linnaeus.
BANISTERIA PRAENUNTIA, new species.
Plate 17, fig. 2.
Description.-Leaves of medium size, broadly ovate in general outS le, with an abruptly acuminate tip and a broad rounded or cuneate Sase. Length about 8 cm. Maximum width, at or slightly above the
~ Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 257, pl. 57, fig. 8; pl. 109, fig. 6,
1916.
S Ettingshausen, C. von, Roy. Soc. London Proc., vol. 30, p. 235, 1880. SU. S. Geol. Survey Prof. Paper 91, p. 257.






36 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

middle, about 5 cm. Margins entire, full and rather evenly rounded. Petiolar character unknown. Midrib of medium size, uncharacteris- h tic. Secondaries thin, seven or eight opposite to alternate pairs diverge from the midrib at regular intervals at angles varying from 450 in the upper part of the leaf to 550 in the basal part; they ascend 1 in slight but subparallel curves increasing in intensity as they proceed toward the margins with which they become subparallel and eventually camptodrome. Tertiaries thin, mostly obsolete. Leaf P substance thin but apparently of a somewhat coriaceous texture.
The present species receives its name from its supposed praenuntial O relationship to the existing Banisteria sinem~arienis De Candolle, a form ranging from the West Indies to Brazil and whose somewhat variable leaves may be exactly matched by the fossil.
The genus contains upward of eighty existing species, mostly climbing shrubs, confined to the American tropics and largely developed in northern South America. Its geological history goes back id to the Lower Eocene, a species having been described by Watelet from the Ypresian of the Paris basin and four homotaxial species, one based on seeds, having been described by the writer from the Wilcox group of the Mississippi embayment in Western Tennessee and Kentucky. Several additional fossil species have been described from the European Tertiary, from all of which the Panama fossil is P conspicuously different, its major differential character being its relatively short and broad outline.
A species based upon fruits has been described by Engelhardt from the Tertiary of Ecuador.
There are 5 species of Banisteria recorded by Hemsley from Central America, 3 of these in Panama, B. billbergiana Beurling on thE seashore of the island of Manzaiillo. Two additional Panama species of Banisteria are referred to the allied genus Heteroptery8 Kunth by Hemsley.
Occurrence.-Culebra formation. West wall of Gaillard Cut below Miraflores locks (collected by M. I. Goldman). Culebra formation (lower). West wall of Canal opposite Culebra Railroad station (Collected by D. F. MacDonald).
Family EUPHORBIACEAE.
Genus HIERONYMIA Allem.
HIERONYMIA LEHMANNI Engelhardt (?).
Plate 16, fig. 3.
Iiecronymia clihmnanni ENGELHARDT, Uber neue Tertiirpfltanzen Siid-Amert
kas. Abh. Senek. Naturf. Gesell.. vol. 19, p. 11, pl. 2, figs. 1, 2, 1895.
Description.-Leaves broadly elliptical or somewhat deltoid anoi inequilateral in outline, with a shortly acuminate tip and broadly
Engflhairdt, H.. fiber neue Tertiirpflan Sid-Amerikas, Abh. Senck. Naturf. Ge sellscl., vol. 19, p. 14. p. 2, fls. 18 19. 1595.





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 37
funded full lower lateral margins and a very wide, somewhat ob. liquely truncated base. Length about 12 cm. Maximum width, in the lower half of the leaf, about 10 cm. Margins entire, full, and rounded. texture thin but coriaceous. Midrib stout, curved, prominent on the lower surface of the leaf. Secondaries stout, 10 or 11 irregularly paced pairs, prominent on the lower surface of the leaf; they diverge fro the midrib at wide angles which become more acute in the apical diart of the leaf, those on the narrower side are more ascending and sewhat straighter than those on the wide side, all are conspicu usly camptodrome at some distance from the margin. Tertiaries

a hin, mostly percurrent. Areolation of small, isodiametric polygonal meshes, well marked on the under side of the leaf. This large leaf is unfortunately represented by fragmentary materal from a single locality in the Caimito formation. In some aspects its characters suggest a broad Ficu, but it seems clearly dentical with the species described by Engelhardt I in 1895 from ,e Tertiary of Ecuador. I have, however, queried the determina,on because of the broken character of the Panama material. In Mhe illustration I have reconstructed a leaf from a combination of e Panama material with the more complete specimens figured
Engelhardt from Ecuador. The two largest fragments from
anama are indicated on the drawing by tinting. It is unfortunate for purposes of correlation that the present determination can not be inclusive, although in view of other similarities shown between ti the Oligocene plants of Panama and those from the Tertiary of Ecuador, I am disposed to regard the present determination as ,fairly satisfactory.
The genus Hieronymia comprises about a dozen existing species of shrubs and trees confined to tropical America and rather widely distributed from Mexico to Brazil as well as in the West Indies.
Occurrence.-Caimito formation, 7 miles northeast of Bejuca (U.S.G.S. station No. 6840). (Collected by D. F. MacDonald.) ft olection.-U. S. National Museum, Cat. No. 35314.

Order SAPINDALES.
Family SAPINDACEAE.
Genus SCHMIDELIA Linnaeus.
SCHMIDELIA BEJUCENSIS, new species.
Plate 17, fig. 4.
Description.-Leaf or leaflet elongate elliptic in outline, inequilateral. Apex and tip equally and bluntly pointed inequilateral. Margins entire. Texture coriaceous. Length about 11 cm. MaxiLUber neue Tertilirpflanzen Siid-Amerikas, vol. 19, p. 11, 1895.





38 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
mum width, midway between the apex and the base, about 4.5 em. Width on one side of the midrib 21.5 mm., on opposite side 24 mm. Petiole missing. Midrib flexuous, stout, and prominent. Secondaries stout, regularly spaced, mostly immersed, about 7 alternate pairs diverge from the midrib at angles of about 500, curving upward a subparallel and camptodrome in the marginal region. Tertiaries mostly obsolete, a few percurrent ones seen. II
This large and striking leaf is referred to the sapindaceous genus Schmidelia, which comprises about 100 existing species of the equatorial regions of both hemispheres with unifoliate or palmately compound leaves. About half of the species are American where they are confined to the Antilles, Central, and tropical South America. They are sometimes referred to the genus Allophylus Linnaeus (as by Radlkofer) and with the exception of this genus all of the members of the tribe Thouinieae are confined to America. Fossil representatives have been unknown except for the petrified wood from the Oligocene of the island of Antigua which Felix described a., Schmideliopsis.D
Occurrence.-Culebra formation. East wall of Gaillard Cut jus north of station 1760 (collected by M. I. Goldman).
Caimito formation, 7 miles northeast of Bejuca (U.S.G.S. 6840) ] Collected by D. F. MacDonald.)
Type.-Cat. No. 35315, U.S.N.M.

Order THYMELEALES.
Family LAURACEAE.
Genus MESPILODAPHNE Nees.
MBSPILODAPHNE CULEBRENSIS, new species.
Plate 17, fig. 3. I
Description.-Leaves lanceolate-falcate in general outline, wit acuminate apex and base. Margins entire. Texture subcoriaceot, Length about 10 cm. Maximum width, in the middle part of thl bc leaf, about 2.5 cm. Petiole missing. Midrib stout, curved, promineb on the under surface of the leaf. Secondaries stout, remote, regu I larly spaced, nine or ten subopposite to alternate pairs, they diver g from the midrib at angles of about 65 degrees and are conspicuousl; camptodrome close to the margins. Tertiaries obscured by the po preservation of the material.
The present species resembles numerous existing and fossil specie 6 of Lauraceae, from all of which, however, it appears distinct. It ibI similar to Mespilodaphne columbiana Berry of the Upper Claibo of the Mississippi embayment, but is a stouter, more falcate, shorter t and less acuminate form.
1 Felix, J., Die fossile H51izer Westindiens, p. 16, pl. 2, figs. 6. 8. 1883.





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 39

The modern species of Mespilodaphne are numerous, inhabiting frica and tropical America, and are often united with Oreodaphne nd Strychnodaphne to form the composite genus Ocotea of Aublet. heir fossil history is almost entirely lost in the multitude of species at have been referred to the form genera Laurus and Laurophyl. Mespilodaphne is abundant and varied throughout the Eocene nd Oligocene of the Mississippi embayment area. Occurrence.-Culebra formation, upper part. East wall of the aillard Cut just north of Canal Zone station 1760. (Collected by I. Goldman )

Order MYRTALES.
Family MYRTACEAE.
Genus CALYPTRANTHES Swartz.
CALYPTRANTHES GATUNENSIS, new species.
Plate 18, fig. 1.
Description.-Leaves broadly oblong-elliptic in general outline, dest in the middle and tapering equally in both directions to the abruptly acute apex and base. Margins entire. Texture subcoria)us. Length between 7 cm. and 8 cm. Maximum width between .5 cm. and 4 cm. Petiole missing. Midrib stout, somewhat curved, rominent on the lower surface of the leaf. Secondaries thin, very umerous, and close set, often inosculating by forking; they diverge rom the midrib at angles averaging about 70 degrees, at intervals f 1 mm. to 3 mm., pursue a but slightly curved outwardly ascending ourse and have their ends united by an acrodrome vein on each dge of the lamina parallel with and from 1 mm. to 2 mm. within e margin. Tertiaries forming open isodiametric polygonal meshes. The present well-marked species closely resembles the only other amed fossil form Calyptranthes eocenica Berry from the lower ocene of the Mississippi embayment (Wilcox Group). It may also compared with the slightly smaller Myrtus rectinervis described Saporta' from the Sannoisian of southeastern France.
The genus Calyptranthes, which is exclusively American in the ting flora, has about seventy species ranging from Mexico and SWest Indies to southern Brazil. There is a strong generic likebetween the leaves of all of the species. Calyptranthe8 zyzygium Candolle may be mentioned, among others, as a form with leaves ost exactly like the fossil. There is also a marked family resem' ance to some of the existing tropical American species of Eugenia, d more especially Myrcia, Myrcia multiflora De Candolle from e Guianas being very similar to the present species.
Saporta, Atudes, vol. 1, p. 251, pl. 11, fig. 5, 1863.






40 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
Hemsley records 7 existing species of Calyptranthe8 from Central America, two of which occur in Panama.
Occurrence.-Gatun formation, Gatun Borrow Pits. (Collec
by M. I. Goldman.)

Family MELASTOMATACEAE.
Genus MELASTOMITES Unger.
MELASTOMITES MICONIOIDES, new species.
Plate 18, fig. 2.
Description.-Leaf oblong-elliptic in outline, of relatively small size, with an equally and bluntly pointed apex and base. Lengt about 6 cm. Maximum width, in the middle part, about 2.25 cr Margins entire. Texture subcoriaceous. Petiole short and stout Midrib stout and prominent. Lateral primaries stout, prominent diverging from the midrib at an acute angle just above the bas and acrodrome. From the disposition of the outwardly directe nervilles from the primaries it is probable that subordinate acrc drome primaries constitute an infra marginal vein on each side, bt these can not be made out. Close-set subparallel nervilles run tram versely between the midrib and the primaries.
This species is represented by a small amount of fragmentary material, too poor to permit definite generic determination. It i therefore, referred to the form-genus Malastomites proposed by U I ger for generically undeterminable leaves of the Melastomatacea While the fossil somewhat suggests the leaves of various Lauraceo genera, such as Cinnamomum, Camphoromaea, Goeppertia, ar ? Cryptocarya, its characters are clearly those of the Melastomataces & It particularly suggests the genus Tibouchina Aublet, which has u, t ward of 200 species of shrubs and undershrubs in tropical Americ '
The family Melastomataceae is a relatively large one, with abo150 genera and over three thousand species. It is almost strict M0 tropical, although some members range southward to 400 south la ( tude. This great family is typically American, seven of the fifte TI tribes into which it is divided being confined to tropical Amerik and about 2,500 of the existing species being also endemic in tf r region. While the geologic history of this vast assemblage of for~ f' is practically unknlmown, there is no evidence to disprove the theo that it, like the allied families Combretaceae and Myrtaceae, had origin in that most prolific region-the American tropics.
The few fossil forms that have been found, including leaves, flo itl ers, and calices, have been referred to the form-genus Melatomi .4 first proposed by Unger. A doubtfully determined species, whi probably belongs to the Lauraceae, has been recorded from the U s






GEOLOGY AND PALEONITOLOGY OF THE CANAL ZONE. 41

er Cretaceous of Westphalia. The only known Eocene species is e well-marked form present in the lower Eocene of the Mississippi nbayment region (Wilcox Group.) Four Oligocene species have een described from Bohemia, Styria, and Egypt; four Miocene pecies from Switzerland, Prussia, and Croatia; and a Pliocene Ipecies from Italy.
i Occurrence.-Culebra formation, upper part. East wall of Gailurd Cut just north of Canal Zone station 1760. (Collected by M. I. oldman.)
Order EBENALES.
Family EBENACEAE.
Genus DISOPYROS Linnaeus.
DIOSPYROS MACDONALDI, new species.
Plate 18, figs. 4-8.

Description.-Globose berry-like fruits of small size and considerble consistency, possibly preserved in an unripe state since the flesh stringy and with a great many tannin cells. The great abundance !f these fruits in the andesitic tuffs makes it seem more probable, however, that they are mature, particularly as some are greatly flatj ied. The numerous elongated pendulous seeds and the amount of vascular fibers in the flesh would tend to prevent much compression ,a certain number of cases. Diameter 12 to 15 mm. Flesh hard, cry tanniferous, and with numerous fibers. Seeds 8 to 10 in number, long, elliptical, compressed, with a hard seed coat. The interior f the seeds is filled with amorphous silica and fails to show any structure. Seeds about 7.5 mm. long, averaging 3 mm. high and 1
to 2 mm. thick, very unequally developed, one to three usually ore or less abortive. Peduncle not preserved, nor do any of the pecimens show the calyx.
These seeds are exceedingly abundant and more or less perfectly ilicified, the flesh being dark brown and the seeds white, making cry striking objects. They are clearly referable to Diospyros and o far as I know represent the only known petrified fruits of this us, although the persistent calices are not uncommon as impresions from the Upper Cretaceous onward. The modern species have om 4 to 12 compressed seeds which tend to become less numerous ith the increase in the fleshy part of the fruit, so that possibly these ore consistent and prevailingly 10-seeded fossil fruits may represent I earlier stage in their evolution, although this seems doubtful ince the calyx of a very large fruited form is known from the Upper ocene of southwestern Texas.






42 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

Diospyros is cosmopolitan in the existing flora with about 18 d, species in the warmer regions of both hemispheres. Mostly Oriental is but not uncommon in the southern United States, Antilles, and frou T Mexico through tropical South America. Upward of 100 fossil spe cies are known ranging in age from the Upper Cretaceous to th I present.
Occurrence.-Section near mouth of Tonosi River, in deposits o I Eocene age (MacDonald).
Type.-Cat. No. 35316, U.S.N.M.

Order RUBIALES.
Family RUBIACEAE.

Genus RONDELETIA Plumier.
RONDELETIA GOLDMANI, new species.
Plate 18, fig. 3.

Description.-Leaves lanceolate in outline, somewhat falcate an inequilateral, with an equally acuminate apex and base. Length b( & tween 12 cm. and 13 cm. Maximum width, midway between the ape 6i and the base, about 3 cm., 13.5 mm. on the concave side and 15.5 mn on the convex side. Margins entire. Texture coriaceous. Petio short and stout, expanded proximad, about 5 mm. long. Midri curved, stout, and prominent. Secondaries thin, numerous, subopp ~ site to alternate, rather regularly spaced; about 15 pairs diverge froth a the midrib at angles of about 450 and ascend in rather flat but regul N and subparallel curves and are camptodrome in the marginal regio !ig Tertiaries obsolete.
This well-marked species is referred to the subfamily Cinchonoides ti and tribe Rondeleticae and seems to indicate an Oligocene species c Riondeletia, a genus of shrubs and trees confined to tropical Ameri and not heretofore found fossil. Rondeletia has about 70 existing sp i cies, a few of which occur in northern South America, but the mi jority are confined to the Antilles (45 species) and Central Americ 4 (24 species).' The present species may be compared with the exis i ing Rondelet;a ra(cmosa Swartz of Jamaica, and with other Antillea and Central American forms. More remote comparisons may l1
made with certain species of Psychotria, as, for example, Psychotnt barbiffIora De Candolle of Brazil, and with the genus Tapiria Ju sieu of the Anacardiaceae, a fossil species of which, Tapiria lance. h lata, has been described by Engelhardt 2 from the Tertiary of Ecut Iv SBritton records 35 species from Cuba. Bull. Torrey Bot. Club, vol. 44, pp. 20-30, 193 SEngelhardt, H., iPher neue Tertilirpflanzen Sild-Amerikas, Abh. Senck. Naturf. (esel vol. 19, p. 15. pl. 9, fig. 4. 1895.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 48

or. Another fossil species somewhat resembling the Panama form s Ginchonidium multinerve described by Ettingshausen 1 from the ertiary of Priesen, Bohemia.
Named in honor of Dr. Marcus I. Goldman, who collected it while Fellow at the Johns Hopkins University.
Occurrence-Gatun formation, Gatun Borrow Pits. (Collected by I. Goldman.)

Genus RUBIACITES Weber.
RUBIACITES IXOREOIDES. new species.
Plate 18, figs. 9-12.

Description.-Fruit bilocular, indehiscent or tardily dehiscent, gneous, capsular-like. Form a prolate spheroid 2.7 cm. long and 2 .M. in diameter. The surface roughened by small tuberculations and its. Walls about 2 mm. thick. Median partition thin. Seeds one n each cell, suspended, elliptical in both transverse and longitudinal etions, compressed along the central partition. Surface striate. ndbsperm not ruminating. One seed is more fully developed than e other. The larger is about 2 cm. long, 1.4 cm. wide and 9 mm. hick.
This well marked -orm is unfortunately represented by but a single pecimen which however shows most of the cavity occupied by the Iit, the two contained seeds partially petrified and the lignified wall nd part of the partition. The accompanying illustrations show the ternal appearance of the fruit (fig. 9) and a side view showing the elative development of the two seeds (fig. 10). Figure 12 shows a ignified end of the fruit with the median partition and figure 11 sa side view with the smaller seed in front and the larger forming he background. So far as I know nothing like it has previously n found fossil.
There seems to be no question but that the present fruit represents one Oligocene species of Rubiaceae and it is consequently referred
the form-genus Rubi cites proposed by Weber, although probably ot congeneric with the previously described fossil species of Rubiates. The fruits of this large family exhibit considerable variety ing either capsular, achene-like or drupaceous. Without a much .rger amount of recent comparative material than is available it is Sot possible to definitely fix the botanical relation of the present pecies which, however, appears to be referable to the tribe Ixoreae or he Psychotrieae. The specific name chosen suggests a resemblance Sthe fruits of Ixora Linnaeus, a genus with over 100 species of
aEttingshausen, C. von, Die Fossile Flora des Tertlitr-Beckens von Bilin, Theil 2, p. 208, 36, fig. 5, 1868.
8370*-18g-Bull. 103 -4






44 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

shrubs and small trees found in the tropics of both hemispheres bu chiefly Asiatic.
Occurrence.-Gatun formation. Gatun Borrow Pits. (Collecte( by M. I. Goldman.)

EXPLANATION OF PLATES.

PLATE 12.
Palmoxylon palmacites (Sprengel) Stenzel. Cucuracha formation.
FIG. 1. Showing abundance of fibrovascular bundles and gum cells. X20.

PLATE 13.
Fio. 1. Ficus ilebrcni.,is Berry. Culebra formation.
2. Guatteria culebrensis Berry. Culebra formation.
3. Myristicophyllumnt panamense Berry. Culebra formation.

PLATE 14.

Taenioxylo imnultiradiatumn Felix. Culebra formation. FIG. 1. Transverse section. X25.
2. Same. X200.
PLATE 15.

Taenioxylon multiradiatum Felix. Culebra formation. FIG. 1. Radial section. X200.
2. Tangential section. X200.

PLATE 16.

FIG. 1. Cassia culebrensis Berry. Culebra formation.
2. Inga oligocaenica Berry. Culebra formation.
3. Hieronymia lelhmanni Engelhardt (?). Caimito formation.

PLATE 17.

Fi. 1. Hiraca oligocaenica Berry. Caimito formation.
2. Banisteria praenuntia Berry. Culebra formation.
3. Mespilodaphne culebrensis Berry. Culebra formation.
4. Schmidelia befucensis Berry. Caimito formation.

PLATE 18.
FIG. 1. Calyptranthes gatunensis Berry. Gatun formation.
2. Melastomites miconioides Berry. Culebra formation.
3. Rondeletia goldmani Berry. Gatun formation.
4-8. Diosypros macdonaldi Berry. Eocene (?).
4. Showing abundance of fruits in tuffs. 5, 7, 8. Transverse median sections of fruits.
6. Longitudinal median section of fruit.
9-12. 1Rubiacites ixoreoides Berry. Gaun formation.
9. External appearance.
10. Median longitudinal section showing unequally developed seeds.
11. Side view of seeds.
12. Llgnified fragment showing end walls and partition.




















S. NATIONAL MUSEUM BULLETIN 103 PL. 12






















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FOSSIL DICOTYLEDONOUS LEAVES. FOR EXPLANATION OF PLATE SEE PAGE 44.







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TAENIOXYLON MULTIRADIATUM FELIX.

FOR EXPLANATION OF PLATE SEE PAGE 44.






E U. S. NATIONAL MUSEUM BULLETIN 103 PL. 16



































































FOSSIL DICOTYLEDONOUS LEAVES.
FOR EXPLANATION OF PLATE SEE PAGE 44.








U. S. NATIONAL MUSEUM BULLETIN 103 PL. Ii


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U. S. NATIONAL MUSEUM BULLETIN 103 PL. 18

























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INDEX.



Page. PAgM
Acratichum ----------------16. 18, 22, 23 Inga latifolia,-----------------------33
aureum ----.-------------23 oligocaenica -------------18, 22,32, 44
kI~hyus------- ------- 38 Ixora-----------------------------4
A nona -------- ----------------19, 28, 29 Laguncularia -------------------glabra ------------- -----------28 Laurus -------------------------- 9
marcgravil --------------------27 Laurophyllum -------------------reticulata---------------------28 Melastomites ------------------------40
Asimina --------------------------28, 29 micouloldes ------18, 23, 40, 44
triloba --------------------28, 29 Melodorum -------------------------28
Avicnnia ------------------------ 16, 22 hMespilodapline --------------------38, 39
Bactris ----------------------------16 columblana,-------------38
Bateria---------------------20, 85, 36 culebrensis ---18, 23, 8, 44
bilbergiana ----------------36 Mitrephora -------------------------28
praenuntia ---------18, 23, 35,44 Myrcla ---------------------------21,p 3,q
sinemariensts --------------36 multiflora --------------------39
Catranthes --------- ---------21, 89, 40 Myrlstica -------------------------20, 30
eocenica -------------------39 Myrlsticophyllum ------------------20,29
gatuneflsl8---------*18, 23, 39, 44 panamense-- 18, 22, 29,44
zyzygiUM ------------------39 Myrocarpus -------------------------34
Oamphoromaea ----------------------40 Myrtus rectinervis -------------------39
C~assa--------------------------- 20, 34 Ocotea -----------------------------39
culebrensis ------------18, 22, 34,44 Oreodapline -------------------------39
ruseifolia.---------------------33 Palmacites dublus -------------------24
Cbaaeorea ------------- -----------16 tenerum ------------------24
Cihnidium multlnerve --------------43 Palmoxylon -------------------17, to, 25
Cnaomum ----------------------40 antiguense ----------------25
Cmsneura -----------------------30 aschersonl,----------------25
Carpus -----------------------16,22 cey'lanicum ---------------25
Crstia --------------------------16 integrum -----------------25
Crytoarya,------------------------40 mississippiensis -----------25
Di1spyros--------------------------- 21, 41 palmacites --------18, 22, 24, 44
macdonaldi ---------18, 23 41,44 stellatum.-----------------25
Enogenites palmactes -- ...---------24 tenerum ------------------25
-------------- --- ---21 variabile -----------------25
F4cultes palmaites--..-..-----------24 Palms ----------------------16, 18, 23, 24
Ficus----------------------------19,26 Parltlum--------------------------16
culebrensis -------------18, 22, 26,44 Psychotria, barbiflora -----------------42
jyn ------------ ----- 27 Rhizophora -----------------------16, 22
newtonensls --------------------26 Rondeletia ------------------------21, 42
Goepertia -------------------------40 goidmanni ---------18, 23, 42,44
Guatteria------------------19,20,279,29 racemosa ------------------42
culebrensis-----18, 22, 27, 44 Rubiacltes ------------------------21, 43
dolichopoda --------- 27 ixoreoldes ----------18, 23, 43, 44
grandiflora 27 Schmidella ---------------------20,37, 3S
ouregon------ -- ----- 27 bejucensls ---------18,23,37,44
Hetropterys--------- ------36 Sophora ----------------------------34
Hfippomane-------- ---16 Swetia ----------------------------34
-------------------20,835 Taenioxylon ----------------17, 20, 30, 32
Chrysophylla ------------------35 multiradlatum --18, 22, 80,44
oligocaenica----------- 18, 23, 35p44 varlans----------------- 31
wilcoxiana --------------------35 Tapiria lanceolata -------------------42
Hieronymia---------17, 20, 36,37 Tibouchlna -------------------------40
Inga lehmanni ---------18,23, 36,44 Toluifera ---------------------------34
-------------------------- 20,329,33 Tritlinax -------------------------16
densiflora ----------------------33 Venericardia planicosta ---------------17
edulis --------------------------33 Virola -----------------------------30







SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM Bulletin 103


CONTRIBUTIONSS TO THE GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE, PANAMA, AND GEOLOGICALLY RELATED AREAS IN CENTRAL AMERICA AND THE WEST INDIES




THE SMALLER FOSSIL FORAMINIFERA

OF THE PANAMA CANAL ZONE


By JOSEPH AUGUSTINE CUSHMAN
Of the United States Geological Survey





Extract from Bulletin 103, pages 45-87, with Plates 19-33

















WASHINGTON
GOVERNMENT PRINTING OFFICE 1918









THE SMALLER FOSSIL FORAMINIFERA OF THE PANAMA CANAL ZONE.


By JosEPH AUGUSTINE CUSHMAN, Of the United States Geological Survey.

INTRODUCTION.

The collection of fossil foraminifera included in this report were sent to the writer by the United States Geological Survey. It consists almost entirely of material collected by Messrs. D. F. MacDonald and T. Wayland Vaughan in 1911, to whom I am indebted for data as to the geological correlation. The names applied to the logic formations are those used in MacDonald's "Sedimentary operations of the Panama Canal Zone, with special reference to the 'stratigraphic relations of the fossiliferous beds," which appears in the latter part of this volume. Where former correlation has seemed not to apply to the foraminifera, especially those of three stations,
60 6035, and 6036a, discussion of the data obtained from the foraminifera is given in detail later.
The orbitoids and nummulites are both well represented in the collection, but as these require special study in connection with those of the Coastal Plain and of the West Indian region it seems advisable to treat them in a separate paper which immediately follows the present one.
The following data are given for only the stations from which foraminifera were obtained and which are recorded in this paper.

LIST OF MATERIAL.
LS.G.S station 6009.-Oligocene-Culebra formation (upper part).
From section in Canal cut 600 feet south of Miraflores Locks.
Dark, soft, fairly well laminated clay rock.
Few foraminifera and rather poorly preserved. W50)-Oligoene-Culebra formation (lower part).
From section-Pedro Miguel Locks to Paraiso Bridge.
Dark, well laminated, very soft, carbonaceous clay rocks.
Foraminifera in fairly good numbers and a rather varied assortment; mostly stained black, except certain of the Miliolidae, which still keep their calcareous tests more or less in their
original condition.
45






46 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

6019 .-Oligocene-Culebra formation.
From section-west side of Gaillard Cut.
a. Dark, well laminated soft and very friable carbonaceous shale
Few foraminifera-some glauconitic, others well preserved.
c. From a lens of sandy limestone 5 feet thick.
Few foraminifera-some stained, some glauconitic, rather poorly
preserved as to details.
d. From lenses of limy sandstone at base of gravel, 3 feet thich
Few foraminifera and these poorly preserved. 6015.-Oligocene-Emperador limestone.
From old quarry, one-fourth mile north of west from Empir
Cream-colored, coral limestone.
Few foraminifera.
6016.-Oligocene-Emperador limestone.
From old quarry, one-third mile north of west of Empire.
Few poorly preserved foraminifera.
6019.-Section on west side of Gaillard Cut near Las Cascadas.
a-f. Oligocene-Culebra formation.
a. Grayish, rather nodular, impure limestone.
Foraminifera few and poor.
b. Dark, well stratified, very friable, tufaceous material.
Foraminifera few and poor except Orbitolites, which are lar
and fine.
c. Grayish, well stratified, very friable, tufaceous sandstone.
Few casts of foraminifera and central portions of orbitoids.
d. Grayish-green, limy, tufaceous sandstone.
Very few foraminifera, poor specimens.
e. Thin-bedded, light gray to cream-colored, limy sandstone wi
some partings of light-colored clay.
orbitoids and Orbitolites? only.
f. Dark, very friable shales and tuffs.
Foraminifera fairly common, some well preserved, others gl:
conitic.
g. Oligocene-Emperador limestone.
Light gray to yellowish gray, somewhat sandy limestone.
Some orbitoids and Orbitolites? but little else in the way
foraminifera.
6020.-Oligocene-Culebra formation.
Same locality as 6019.
a-c. Dark-gray carbonaceous clays, friable shales and tuffs.
a. Foraminifera numerous but of few species, mostly glaucon
at least in part.
c. A few Orbitolites in the coralliferous layer.






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 47

O04.-Section in railway cuts near New Frijoles.
a. Oligocene-Culebra formation.
Dark, basic, orbitoidal, tufaceous material.
Many worn central portions of Orbitoids and a very few other
foraminifera poorly preserved.
5.-Oligocene-Culebra formation (upper part). About 200 yards south of southern end of switch at Bohio
Ridge station relocated line Panama Railroad.
Contains a number of species of foraminifera but for the most
part broken or poorly preserved. 026.-Two miles south of Monte Lirio.
Somewhat coarse-grained sandstone.
Few poor specimens of foraminifera.
19.-Setion one-half mile from Camp Cotton, toward Mionte Lirio, at big curve on railroad. Miocene-Gatun formation. a. Bluish, fossiliferous argillite.
Very few foraminifera.
b. Bluish argillite.
Few foraminifera, but considerably more than in a.
c. Bluish, fossiliferous argillite.
Very few poor specimens of Amph!stegina.
0.-Railroad cut north side of Big Swamp, one and ofe-half miles north of Monte Lirio. Miocene-Gatun formation.
Bluish gray, argillaceous beds.
The only foraminifera consisted of a single specimen of Trilo.
oulina.
'01.-Section in cut one-half mile west of Camp Cotton toward Gatun. Miocene-Gatun formation.
Conglomerate bed and sandy marl 1 foot above.
A few poorly preserved specimens of Quinqueloculina were the
only foraminifera.
053.--Generalized section of the bluffs exposed along the Panama Railroad, relocated line, about 3,500 feet south of Gatun Railroad Station. Miocene Gatun-formation.
c. Dark-colored, marly, fossiliferous clay.
Rich in foraminifera, especially in specimens. A fair number
of species, well preserved.
0'5.-Vicinity of Mindi Hill. Miocene-Gatun formation.
Gray-green, fine grained sandy shell marl.
Very fine-grained material, but with numerous species and specimens of foraminifera representing an off-shore assemblage.
086.-Monkey Hill, Mount Hope Station. Miocene-Gatun formoation.
Dark-colored, fine grained, sandy clay marl.





48 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.

Specimens of foraminifera numerous and well preserved, re]
resenting an off-shore assemblage comparable to 6035.
5850.-Near Mount Hope-Pleistocene.
Loose shells and marl obtained from ditch through swami'
ground about one-fourth mile from present sea beach aIl
about 6 to 8 feet above high tide.
Contains a few foraminifera of common shallow water, tropic
species.
The geological position of certain material from near the Atlan1 end of the canal seems from the evidence of the contained forami, fera to be younger than the position previously assigned to it-t'i upper Oligocene. By a reference to the table of distribution it w be noted that the great majority of the species occurring at the st tions in question; 6533c, 6035, and 6036, do not occur in the rn terial of definitely Oligocene age. In such cases as that of Grist laria rotulata there is a slight difference in the specimens from tht stations and those from the Pacific side, 6010, 6012a, 6012c, but t specimens at the latter stations were in small quantity, and the d ferences could not be made use of, mainly from lack of a sufficie number of specimens. In the case of Cristellaria vaughani tl' seems to be a well-characterized species occurring at several statioi but even in it there are very minor differences. Among the spec i of Globigerina, the more generalized species such as G. bulloid which has a very wide geological range, occur more or less constant throughout the collections, but the strongest evidence comes fr( the last three species and Orbulina, which are very rarely fou fossil, and then only in the very latest tertiary. These were w characterized species, the specimens are very clean and complete, a resemble a modern Globigerina ooze of considerable depth. q three species of Pulvinulina also occur nowhere but at these station Pulvinulina concentrica is essentially a recent species and P. T nardii is characteristic of modern Globigerina ooze. Sigmoilo tenus and S. asperula are also speces of recent Globigerina ooze moderate depths. On the other hand, the lack of certain things also significant. Amphistegina, which occurs more or less regula in the other portion of the material, is entirely wanting in the th Pacific stations, 6033c, 6035, and 6036. Polystomella als does occur. Both the last two genera are very characteristic of coastal plain Oligocene of the United States. It may be argued this case, however, that the stations were originally too far fr shore to have these genera which are more characteristic of shall littoral conditions.
On the whole, the foraminifera bear out the geological determi' tions based upon the other groups of organisms.









GEOLOGY AND) PALEONTOLOGY OF THE CANAL ZONE. 49







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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 51
DESCRIPTIONS OF SPECIES.

Family TEXTULARIIDAE.

Genus TEXTULARIA Defrance, 1824.

TEXTULARIA ABBREVIATA d'Orbigny.

Plate 19, fig. 1.

Textularia abbreviata D'ORBIGNY, Foram. Foss. Bass. Tert. Vienne, 1846,
p. 249, pl. 15, figs. 9-12 (7-12).
Description.-Test broad and short, somewhat compressed, chainers comparatively few in number, broad near the center and taperng to the periphery, sutures in these specimens indistinct, aperture n arched slit extending nearly across the test, wall comparatively mooth.
Length 0.65 mm., breadth about 1 mm. Cat. No. 324608, U.S.N.M.
Specimens from U.S.G.S. No. 6010, from the Culebra formation, ark clay north of Pedro Miguel Locks. Apparently the material Rather metamorphosed and more or less glauconitic so that little f the original test is preserved. This is a rather common Tertiary species.
TEXTULARIA SAGITTULA Defrance.

Plate 19, fig. 2.

Tetulari a sagittula DEFRANCE, Dict. Sci. Nat., vol. 32, 1824, p. 177; vol. 53,
1828, p. 344; Atlas, Conch., pl. 13, fig. 5.
Description.-Test. elongate, tapering, much compressed especially A sides, chambers numerous, sutures indistinct, aperture a curved 31it occupying about one-half the width of the base of the chamber.
Length about 1.5 mm., breadth 1 mm. Cat. No. 324609, U.S.N.M.
A few poorly preserved specimens from U.S.G.S. No. 6025, from he Culebra formation, foraminiferal marl and coarse sandstone bout 200 yards south of southern end of switch at Bohio Ridge tation, relocated line, Panama Railroad.
Although this material is more or less glauconitic and poorly prerved the three specimens, one of which is here figured, are referred vith a reasonable degree of certainty to this species. A single fragmentary, specimen from U.S.G.S. No. 6026, from the J1ebra formation, coarse, sandy foraminiferal marl about half way Between Monte Lirio and Bohio Ridge, relocated line, Panama Raiload, seems also to be this species.



k|





52 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
TEXTULARIA AGGLUTINANS d'Orbigny.

Plate 19, fig. 3.

Te.xtuaria agglutinans D'ORBIGNY, in De la Sagra, Hist. Fis. Pol. Nat
Cuba, 1839. Forarminif Description.-Test elongate, tapering, but slightly compressed lat-~ erally, chambers high, sutures deep, outline sinuous, end view broadly elliptical, wall composed of rather coarse agglutinated material, aperture a narrow slit a little more than half the width of the base of the chamber.
Length 1.23 mm., breadth 0.65 mm. Cat. No. 324610, U.S.N.M.
A single specimen here figured seems referable to this species. Ii is from U.S.G.S. No. 6019-f, from the uppermost bed of the Culebre formation, the lower limestone of the Las Cascadas section, opposit Las Cascadas, Gaillard Cut. Although not so rounded in end viev as this species usually is in recent specimens, the general characterE wall structure, high rotund chambers and lobulated outline seem t'f place it here.
TEXTULARIA LAMINATA, new species.
Plate 19, fig. 4.
Description.-Test elongate, cuneate, tapering from the wide
part near the apertural end, gradually and evenly to the initial en which is subacute, median line raised thence tapering rapidly towar the periphery which is thin and extends out into a lamella-lilk border, chambers numerous, wide and low, sutural lines raised, som what curved backward; border irregular, wall finely arenaceowi aperture indistinct.
Length 2 mm., breadth 1.2 mm.
Specimen figured from U.S.G.S. No. 6010, from lower part of tl Culebra formation, dark clay north of Pedro Miguel Locks. Spec men rather better preserved than most from this station. The er view of this specimen is mainly rhomboidal with the borderir carina extending outward in a thin carina. It is in some ways su festive of Textularia carinata but differs in many respects fro that species which is also figured on plate 19, fig. 6.
Type-.specimen.-Cat. No. 324611, U.S.N.M.

TEXTULARIA SUBAGGLUTINANS, new species.

Plate 19, fig. 5.

De(rip;;tion.--T5est sulrhomboidal in front view tapering from t. middle toward either end, in end view oblong, sides truncated; cha Lers comn)aratively few, somewhat inflated, sutures conspicuously <






GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 53

ressed, wall composed of rather coarse arenaceous material, aperure extending into the base of the chamber in a narrow rounded pening deeper than wide.
Length 1.3 mm., breadth 0.85 mm. This species was fairly common from U.S.G.S. No. 6033c, the atun formation, in marl from second bed from bottom, just below ower clay, Gatun section, relocated line Panama Railroad. This species may be distinguished from Textularia aggutinans by he truncated sides, the oblong end view and especially by the deep, iirrow aperture.
Type specimen.-Cat. No. 3'4612. U.S.N.M.

TEXTULARIA CARINATA d'Orbigny.

Plate 19. fig. 6.
Te.rtularia carinlul D'Im v. Ann. Sci. Nat.. vol. 7. 1826. p. 263, No. 23:
Foram Foss. Bass. Tert. Vienne, 1846, p. 247, pl. 14, figs. 32-34.
Descrption.-Test much compressed, rather abruptly tapering oward the initial end, sutures strongly limbate, in well-preserved pecimens extending out from the periphery in angular spine-like rojections, aperture narrow. elongate. Length 1 mm., breadth 0.65 mm. Cat. No. 324613, U.S.N.M.
The only minaterial..of this species is from U.S.G.S. No. 6036, from he Gatun formation, a dark-colored, fine-grained, sandy clay marl om Monkey Hill, Mount Hope Station. It is very evidently this pecies and is well preserved.

TEXTULARIA PANAMENSIS, new species.

Plate 20. fig. 1.
Description.-Test rhomboid in front view, very much compressed, Send view long and narrow, the faces nearly parallel, sides rounded: Composed of comparatively few chambers but variable; long and ow, sutures somewhat depressed, wall rather coarsely arenaceous; perture indistinct.
Length 0.85 mm., breadth 0.65 mm. The figured specimen is from U.S.G.S. No. 6036, from the Gatun .ormation, a dark-colored, fine-grained sandy clay marl from Monkey ill, Mount Hope Station. Specimens were common from U.S.G.S. 10. 4-033c, in marl from second bed from bottom, just below lower lay, Gatun section, relocated Panama Railroad. This is a rather striking species, with its very flat, broad front iew and very compressed character of the test. Type-specime.-Cat. No. 324614., U.S.N.M.






54 BULLETIN 103, UNITED STATES NATIONAL MUSEUM.
Genus CHRYSALIDINA d'Orbigny, 1846.
CHRYSALIDINA PULCHELLA, new species.
Plate 20, fig. 2.
Description.-Test elongate, gently tapering, broadest at the apica end; in end view triangular; early chambers triserial, later ones uni serial; chambers in uniserial portion triangular, the sutures distinct gently curved backward at the angles, outline more or less irregula apertural face gently convex, with indications of numerous circula apertural openings, wall smooth.
Length 0.5 mm., breadth 0.2 mm.
This species occurred at U.S.G.S. No. 6036, the Gatun formatiorI in dark-colored, fine-grained, sandy clay marl, from Monkey Hil Mount Hope Station.
The species differs from the only known recent species, Chrys6 lidina dimorpha, in the more tapering and elongate test, the greate irregularity of the contour and test in general and its generally let, trim and neat appearance. The specimen figured is well preserve in its general characters, except those of the apertural face, whic are somewhat obscured.
Type-specimen.-Cat. No. 324615, U.S.N.M.
Genus BOLIVINA d'Orbigny, 1826. BOLIVINA ef. B. PUNCTATA d'Orbigny.
Plate 21, fig. 3.
Bolivina punctata D'ORBIGNY, Voyage Ambnr. Mrid., vol. 5, pt. 5, "Foran
niferes," 1839, p. 63, pl. 8, figs. 10-12.-H. B. BRADY, Rep. Voy. C7h lenger, Zoology, vol. 9, 1884, p. 417, pl. 52, figs. 18, 19.-FLINT, Ar
Rep. U. S. Nat. Mus., 1897 (1899), p. 292, pl. 38, fig. 1.
Description.-Test much elongate, sides nearly parallel, abrupt tapering at the initial end, chambers numerous, usually higher thi broad, inflated, sutures distinct but slightly depressed; wall fine punctate, occasionally becoming slightly striate.
Length 0.60 mm., breadth 0.15 mm. Cat. No. 324616a, b, U.S.N.] Specimens which seem referable to this species were obtained U.S.G.S. No. 6033c, Gatun formation, marl from second bed frc bottom, just below lower clay, Gatun section, relocated line Panar Railroad and 6035, Gatun formation, from gray green, fine grain sandy shell marl, vicinity of M indi Hill. There is a tendency f the specimens to take on a semi-striate appearance, an extreme foi both in shape and striation shown in plate 21, figure 3.
BOLIVINA AENARIENSIS (Costa).
Plate 21, fig. 2.
Bricliiw anariensis COSTA, Atti Acad. Pontaniana, vol. 7, 1856, p. 297,
15, fig. 1, A. B.
Bolivina aenariensis H. B. BRADY, Proc. Roy. Soc. Edinburgh, vol. 11, 1
p. 711; Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 423, pl. 53, f
10, 11.





GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 55

Description .-Test much compressed, composed of numerous chainrs about twice as broad as high, sutures distinct, slightly curved backward, chambers slightly inflated, especially in the center, test ordered by a narrow but distinct carina; surface smooth except for ,veral longitudinal raised costae radiating from the initial end which
carries also a short spine.
Length 0.65 mm., breadth 0.35 m. Cat. No. 324617a, b, U.S.N.M.
A few specimens were obtained from U.S.G.S. No. 6033c, Gatun
ormation, in marl from second bed from bottom, just below lower
-lay, Gatun section, relocated line, Panama Railroad. While these specimens are not absolutely typical they undoubtedly
long to this species.
Very typical specimens occur at U.S.G.S. No. 6036, Gatun formaion, in dark colored, fine grained, sandy clay marl, from Monkey
s Mount Hope Station.
Al
BOLlVVNA ROBUSTA H. B. Brady.

Plate 21, fig. 4.
Bolivina robusta H. B. BRADY, Quart. Journ. Mier. Sci., vol. 21, 1881, p.
57; Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 421, pl. 53, figs. 7-9.
DWescription.-Test compressed, gradually tapering toward the
pical end; chambers comparatively few; about twice as broad as igh; sutures limbate, gently curved backward, often slightly lobuated or occasionally showing traces of reticulation on the surface,
all otherwise smooth but punctate, not spinose at the apical end.
Length 0.45 mm., breadth 0.25 mm. Cat. No. 324618, U.S.N.M.
These specimens, an extreme form of which is figured, are many of '~ fhem very close to typical B. robusta which is at best either a variable
species or one including more than one form. The sutures are
ally liimbate, as shown in some of Brady's figures, but no apical Upine is apparently in any of the specimens in this material. They ere from U.S.G.S. No. 6035, Gatun formation, from gray green,
e pained, sandy shell marl, vicinity of Mmdi Hill.

BOLIVINA, species?
Plate 21, fig. 1.
This specimen is rather ill-defined and cannot be definitely determed from the single example, the sutures are limbate as in Bo,i robusta Brady, but have apparently no secondary extensions S in that species. The whole specimen seems to be replaced. The Scimen is from U.S.G.S. 6010, lower part of the Culebra formation,
m dark clay north of Pedro Miguel Locks. Cat. No. 324619,






56 BULLETIN 103., UNITED STATES NATIONAL MUSEUM.

Genus BIGENERINA d'Orbigny, 1826.
BIGENERINA NODOSARIA d'Orbigny.
Platp 21, fig. 5.
Bigenerina nodosaria D'ORBIGNY, Ann. Sci. Nat., vol. 7, 1826, p. 261, I
11, figs. 9-11.-H. B. BRADY, Rep. Voy. Challenger, Zoology, vol.
1884, p. 369, pl. 44, figs. 14-18.
Description.-Test elongate, subeylindrical, early portion consis ing of a few chambers arranged as in Textularia, later ones uniserio early portion tapering abruptly toward the apical end, wall coarse arenaceous, sutures rather indistinct, aperture circular and central.
Length 2 mm., breadth 0.8 mm. Cat. No. 324620, U.S.N.M.
Several specimens in excellent condition were obtained fro U.S.G.S. No. 6036, Gatun formation, in dark-colored, fine-graine sandy clay marl from Monkey Hill, Mount Hope Station.
These specimens, as in the one figured, have but a slight indication of the biserial chambers from the exterior, but otherwise seem to e typical. At first glance they might be taken for a species r Clavulina.
Genus GAUDRYINA d'Orbigny, 1839.
GAUDRYINA FLINTII Cushmnan.
Plate 20, fig. 4.
Gaudryina subrotwndata FLINT (not G. subrotundata Schwager. 186
Ann. Rep. U. S. Nat. Mus., 1897 (1899), p. 287, pl. 33, fig. 1.
G tudryina flintii CUs u~LN, Bull. 71. UIT. S. Nat. Mus., pt. 2, 1911, p. 1,
fig. 102a-c.
Description.-Test subconical, early portion rounded conical, tri r rial, later portion subcylindrical, biserial chambers of later port : nearly semicircular in transverse section, sutures distinct; wall a naceous; aperture subcircular, at the base of the inner margin of -e chamber.
Length 1.20 mm., breadth 0.72 mm. Cat. No. 324621.
A single specimen which seems to be close to recent specimens this species was obtained from U.S.G.S. No. 6010, lower part of e Culebra formation, in dark clay, north of Pedro Miguel Locks. 'e specimen is somewhat glauconitic and certain of the details are me or less obscured.

GAUDRYINA TRIANGULARIS Cushman.
Plate 20, fig. 3.
Gaudryina triangularis CUSnMAN, Bull. 71, U. S. Nat. Mus., pt. 2, 16 1
p. 65, figs. 104a-c.
Description.-Test somewhat longer than broad, early portion angular, the faces somewhat concave, triserial; later portion bisesl