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ANNALS

OF THE

NEW YORK

ACADEMY OF SCIENCES

VOLUME XxXVI

1915

Editor

ae ’ EDMUND OTIS HOVEY

ES ya New York

| Ee Published by the Academy 235 TOS

1915, 1916

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum oF Naturat History, 1817-1876)

OFFICERS, 1915

President—GerorceE F. Kunz

Vice-Presidents—CuaRLES P. BERKEY, RayMOND C. OSBURN,

CHARLES BASKERVILLE, CLARK WISSLER

Corresponding Secretary—Hernry E. Crampton, American Museum

Recording Secretary—Epmunpv Otis Hovey, American Museum

Treasurer—HENnyY J. CocHrRAN, 389 Fifth Avenue

Librarian—Ratru W. Tower, American Museum

Editor—Epmunp Otis Hovey, American Museum

SECTION OF GEOLOGY AND MINERALOGY an

Chatrman—Cuar.es P. BerKey, Columbia University

Secretary—Dovucias W. JoHNnson, Columbia University

SECTION OF BIOLOGY :

Chairman—RayMonvd C. OsBurn, Connec ticut College, New London

Secretary—Witi1am K. Gregory, American Museum |

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY , {4

Chairman—CuHar es BAsKERVILLE, College of the City of New York

_Secretary—Ernest E. Surru, 50 Hast 41st Street

i ard ¥ 7

Ae fy

CONTENTS OF VOLUME XXVI

Page

PT EME Tice noises «6 cisleare sb eaves witcha ina PEt oe eh tars eters asia Sia ra eke ehkte i

Abe RIE yal creer aaah 06 foo ue ahen sie ee anere Lica ca djlehareterale Seve le 8 vac vw o'e-eha ete © ii

SNR MRT oes A Seas has Spe ah scant cs 3 oho c a oe wile lei atalenk clap aie ate a tobereieie: a ld bee be ede'e ili

Mates of publication and editions of the brochures.............0....00.00- iii

ee MN STEER VOLTS 289 che i 7os n> oo, wie eek fenahover Bier see.o, aneiei ah aval Oe enw, Sealed eves od) eles iv

Geological Reconnoissance of Porto Rico. By CHARLES P. BERKEY.

re He Spee DUNN Recerca ad cl.’ SPais are gatebentrratte, Sea ooal'ah st asta Si ahaP ONG Le cen area eevee wie 1

List of Greater Antillean Spiders, with Notes on Their Distribution.

Oe ee SE GU ET Actor Vais. ilsie's, OR Meebel sya Where abla Ge eels s OEMS ese « 71

Volcanic Dust Veils and Climatic Variations. By HENrykK ARCcTOWSKI... 149

Statistical Study of Variation in Spirifer mucronatus. By CHARLES C.

eGR rc ee ia ate ants ol aecuase’ os wha tee) « MUPRTe a rOhah Re Ray GAL: of & IO ws Ra AAW oats 175

Review of the Pleistocene of Europe, Asia and Northern Africa. By

Pasa ATRETE ED OSBORN cairo sais wo o.0 che bo bi detea ward ne rine here kere 215

Present Status of the Problem of the Origin of the Tetrapoda. By Wi-

Dee GHCoOny. CP late DV )iic. occas & od sccam clea eo sce clen Mee ore Siete ‘317

Biochemical Studies of Selenium. By Victor E. Levine. (Plate V)..... 385

igecords of Meetings, 1915. By Henry EX. CRAMPTON...........062c-c006 395

Peet tshe ot Mecemmber, 1915.0. 2c. eee oe ae bo eee cee ceee eee 463

a aR ne a Sc aS ee ke eh Shara Qiev dw win bie Gi bhi tebe algls G Suche ab ere iele's 475

DATES OF PUBLICATION AND EDITIONS OF THE BROCHURES

Edition

Pp. 1-70, 3 March, 1915 - 1450 copies

Pp. 71-148, 29 May, 1915 1075 copies.

Pp. 149-174, 21 June, 1915 1200 copies.

Pp. 175-214, 30 June, 1915 1150 copies.

Pp. 215-315, 30 July, 1915 1350 copies

Pp. 317-383, 7 July, 1915 1250 copies

Pp. 385-394, 18 October, 1915 ~1310 copies

Pp. 395-486, 12 May, 1916 1000 copies

iii

LIST OF ILLUSTRATIONS

Plates

I.—Relief Map of Porto Rico.

II.—Geological Reconnoissance Map of Porto Rico.

III.—Generalized Geological Cross-sections of the Island of Porto Rico.

1V.—Right Pectoral Girdle and Limb of Sauripterus taylori Hall.

V.—lIllustrations of Biological Reductions of Selenium Compounds.

Text Figures

Page

Relief features characteristic of the interior ranges of Porto Rico........ 8

Partially destroyed dune sand deposits of the San Juan formation....... 11

Unconformity below the Arecibo formation................. «2. see 16

Typical shale occurrence... . 2. 2 -sie. soc ce « os liaise = hee eles 5 ee 18

Interbedded coamo limestone layers with massive tuffs................. 20

Diorite porphyry sills...... 2.6 c 006s ews See ees ce we be te Cee a ene ee aT.

Photomicrograph of a typical thoroughly indurated dese ash: . s353a8 30

Photomicrograph of a typical weathered shale from Fajardo Playa...... SL

Photomicrograph of a foraminiferal shale from the Bayamon-Comerio

TOBE foc soeeicie a 08 os =m dele wuclon a wie tthe aie sinh wml Oe at a err So -

Photomicrograph of the San Juan formation indurated dune sand....... 33

Shales and ash beds cut by a large irregular dike and sill............... 36

Overturned fold and crush zone in finely bedded shales on the Jayuya

road near the summit of the range......:.......cce..:55 se 38

Crumpled shales as seen along the Jayuya road near the summit of the

TARO 202. ce eo a ele dic eS we de )ataje yes wm delet a Soul Sm ei Sree 40

Cliff of the San Juan formation south of the lighthouse at Arecibo....... 44

Detail of the double structure in the San Juan formation of Arecibo..... 46

Playa plain and marginal ‘terrace.......2..0.c..-+-..-0.0+- 0.0 48

Structure beneath the marginal terraces..................0.20e==nee 49

Inner lowland near Bayamon...........-..20-<ceceeecus css Soe 50

Haystack (pepind) HDills:.......2.0...8. 20...0.2.42..,.02 3 Pry

Cave structure in the haystack (pepino) hills................33 52

View of.a portion of Mona......... 25.0.2... .6+.262...5. 75

View of a portion of Desecheo....... 2.2.6.0. 00500. e even cos TA

sandy plain south of Pifiar del Rio, Cuba..................=3 333 125

View near Bafios San Vincente, Cuba..................2<.+s0 een 126

Distribution of Archwid®..........0....620 04 chen eeecedests 136

Distribution of certain genera known from Baltic amber................ 137

Distribution of Segestriimse. ......0.....000 0c cece cece eccce sl) ae 138

One of the hypothetical upper Cretaceous land masses................-.. 139

Curve of the consecutive means of temperature observed on the summit

of Pike's (Peak... 25 os cae ke Des vc se tabs oe nls cee oe eee 156

Thermopleionian variation of Port Darwin.....................000ne . 158

Temperatures recorded at some Asiatic stations....................ee.. 161

, ate

Text Figures

Page

Wempesimre records at Fort-de-France............6.c00ccenceceereseree 162

Comparison of the consecutive temperature curves of Cayenne and Para

sores ihe West Indteset ts. toemes stn sc... oe alee emis 165

meeuuipa plcion of 1911-1912 observed im Alaska..............00 2. seen... 167

World’s temperature and the variation of the frequency of volcanic erup-

area a See are SEL ake Sew ee Mer ore) ble. aw Saran Me peeyeee aie 173

Curves showing comparative conditions of shell indices in adult and neanic

stages of Spirifer mucronatus mut. alpenense...........ee eee eeeees 178

Curves showing comparative conditions of shell indices in adult and neanic

stages of Spirifer mucronatus mut. multiplicatus......... eee eee eens 179

Curves showing comparative conditions of shell indices in adult and neanic

Stages of Spirifer mucronatus mut. profundus....... 2.20. cceeeseee- 181

Curves showing comparative conditions of shell indices in adult and neanic

stages of Spirifer mucronatus mut. thedfordense...... 0.0... cece eee 183

Curves showing comparative conditions of shell indices in adult and neanic

Stages of Spirifer mucronatus mut. attenuatus......... 2... cece eee. 185

Diagram showing the relationships of the five mutations of Spirifer mucro-

a ease 2s tan taie, hDtn kn StaMe Na lotete te ce Ole Sin /ei ews esl biel sim wie eebs 187

Curves showing comparative conditions of shell indices of the adult stages

BeniveaetatIons Of Spirifer MUCTONATUS « «2 22s wc cc eae ti wees acess 188

Curves showing comparative conditions of shell indices in neanic stages

pepe anmEALIONS OF Spirifer IMUCTONEGTUS .. = 6 os enw oe i wee 189

aaa SO PIE SE EES SSS ov 5. oa Vs vw Shaiya © Cokleie eal, Beis Gin we sins 4 nm bbe sy etatee 224

Skeleton of the Pleistocene pigmy hippopotamus of Madagascar. Hippo-

potamus madagascariensis, together with a skull of the recent hippo-

aN PENILE RITE EID Si 8 on oe at Ss Si Cale Sara sue a vue MLOlS 0 Vd OL SRS we eS 228

Glacial map of northern Germany and the Netherlands.................. 230

Theoretic snow levels during the Glacial Epoch....................-...-.- 231.

Divisions and contemporaneous events of the Glacial Epoch............. 233

Five chief zodgeographic regions of Europe. Asia and northern Africa

from which the mammals migrated into western Europe during the

TTS DP PELL E be) Pie peg BER PRE ER eer eens 2 car oe ene eee 243

Introduction, succession and extinction of the faunz from the five chief

Pe ee OEE MOS nei cps oie mc. a Sala Pecaia a ch lb wb lecteneBarane ele macs Ca shee 247

meerters of North American glaciation... ....... 00560. ewe eee eee eee 253

Principal mammal deposits and culture stations of the Pleistocene of

LS a ee ee ah Nb Nes ee EY IS eite CA SOM ON Sil ete es Chee me eileen o 254

Giant deer, Megaceros, of the British Pleistocene...................00.0% 258

Preeremernereispocene elephants... 22... 00 6. i ae ee ce ewe ew cence 261

Skeleton of Elephas meridionalis of Durfort....................2.-+-2+. 262

Mobtme, Paleolithic and Neolithic implements.................2.20c00:. 265

PeenMieetE Maier near EHeidelbers: .. 22... ec dc ee ve cc b ee we ete ewes 274

eS ete EY De ed a gS osc a ie vid wan ssw cha wna. da'e Sods ola Rw mile Sa eb 275

RE MM RMN ete ogee Beaten CAs Cae ware Sik ota Ok vin veo, 40K 5 ee Wel eles 295

IE NRE Tes a rE re ets Loh, Se eiO ait cits Sa Sits wales dsu as Sembee 296

The hairy mammoth (Elephas primigenius) and Paleolithic man (Homo

Re EEIE SIGE eget hoe uly Ri = Ct Sled: Mua wie a's elute ee hc = da bea'y 6 3 38 bw ee 305

Meeeweiiy rhinoceros (Diceros antiquitatis) ...... 05. ec ec ee cee 306

Text Figures

Page

Rhinoceros skulls. ...- 5 s:cesiveeeGee «+ e+ alleles Deeps etwle elias oa) 307

Pattern of skull-top of: Devonian dipnoans...............-+...---539see 323

Skull pattern of Osteolepis -microlepidotus..........0:¢+.++--+-05meeeeee 330

Skull patterns of Trimerorhachis and Diplopterus............0..0+e000- 336

Pectoral girdle of Acipenser and Amia..... 2.0.0. 00. 520.00. +) eee 347

Pectoral girdle and fin of Ceratodus forsteri...............6..--00:- po 352

Pectoral girdle of Polyterus bichir ... 52% 2.05 66 05 00 010s aimee = 354

Restoration of Megalichthys. After E. D. Wellburn.................... 356

Right pectoral limb of Husthenopteron foordi. After Patten............ 359

Right pectoral limb of Sauripterus taylori—Restoration, medial aspect... 360

Head and pectoral limb of Osteolepis microlepidotus...............002- - 361

Comparison of the paired limbs of Palzozoic rhizodonts and tetrapods... 363

Homology of the coracoid in primitive mammals......................-. 370

Homology of the coracoids in Monotremes.........:.....-.0. see eee 371

Homologies of the coracoids, etc., in reptiles. ........... 2.) 2. see eee 372

Homologies of the coracoids, etc., in Amphibia.......... 0. 42..0e eee 374

a4 So ‘

eS boar eae

: \ Z . C

ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXVI, pp. 1-70, pll. I-III

Editor, Epmunp Oris Hovey :

GEOLOGICAL RECONNOISSANCE OF

PORTO RICO

CHARLES P. BERKEY

NEW: YORK

PUBLISHED BY THE ACADEMY

* a MARCH, 1915

(HE NEW YORK ACADEMY OF SCIENCES

(LiycEUM oF NaTuURAL Hisrory, 1817-1876)

President—Gnorer Freperick Kunz, 601 West ‘Ltoth dizeak -

Vice-Presidents—Cuanres P. Berney, RayMonp C. OsBuRN, Sm

: CTrARLES BASKERVILLE, CLARK WISSLER s

aie est ad pect ela ees Ki. Ces American )

Treasur or—Hnwy HIS aeatiy 389 Fifth cai |

Librarian—Rawteit W. Tower, American Museum — scree

Editor-——KpMUND OTIS Hovey, American Musenm — oo

SROTION OF GBOLOGY in |

Chairman—Crrarces P. ‘Berkey, -Calnathnn University :

Secretary—A. B. Pacint, 2704 Bedford Zak ee ;

SECTION OF BIOLOGY :

Chairman—Ravatond Cc. OsBURN, ‘BBY West 124th ay ee

ee ae cK: oe ke: American Museum

SECTION OF ASTRONOMY, PHYSICS wn ¢

Chairman—CHARLes Reece College of. the on

Secreta 'y—ERNEST Ki. SMuTH 50 East Alst Street

SECTION OF ANTHROPOLOGY AND rs

Chair man—C!LARK W ISSLER, American: ‘Miisetion.

Secr elary— RoBERre H. Lowin, ‘American Museum

‘The sessions of he Perera are held on, ee : rif

o'clock from October to we mon at the ae }

[ANNALS N. Y. Acap. Sci., Vol. XXVI, pp. 1-70, Pll. I-III. 3 March, 1915]

GEOLOGICAL RECONNOISSANCE OF PORTO RICO 1

By CuHartes P. BERKEY |

(Presented wm abstract before the Academy, 7 December; 1914)

CONTENTS

Page

NINE DUEUE REP ce Stara chs he os Siglo oie Cave Soa euler oye One Wes cd mek hoe ee 2

Pee en ToBI CRT fi) avr ate oa eiace so coave wo hc bee REY oe Bet eee 53

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caw BLP ELT BS oS Be? i a Deere Chae Soe ee aM

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PE ete eMNIESL OME Oe occ rl gidc Ps one ede SS RS ek he be US PA

ORME Jae LT HIRE URINE 2 es. os phe wos Sele vw PRS See a tae 22

SAG MEME GLONE Min 2 0 oP oe kat ee fee oO Oe eS 23

eTOCs oo. oer Gia igh Ne Ae ake a Se 23

PRM EEE EGCG cca a2 elas kere Sa wih Fe woul gi eee ahs Sate aS ok De 23

| EES te ORL eee a ee we 25

Ee OS. a eles uss oe cae Ais takes weno ee a eS 26

MIRC INES ih athe ae Skee Soe pe et Shed eae 26

LER TEES SR SE Ie a a a ee a a eR AS 29

Se MIMENEEID ATIC oS 2 ata). i Pace ds kd hv cw ic tlhe nd bok cc oa Om 29

ee PCIE CARE Ba ha a are edi Sl xo hw Beaks EEL Ee 34

Bemmermieeh TCALITCS 66.5 6. ek cece cece cee cues ee tty Seon Ne a SNM are es 35

een RPI tina WES aio. es 5 = ciate ae be oe ee bck bo the bck eae eS 36

Mba SNISDS, FCOUNILEMOS sais) Sa obs od Ged2s bh hace oe loc ek: 37

Les a Ore A ta ae Se a ee en 38

OTT SB IO AEE Se er eee Ne Sc eae ae 39

Seen COheDET el ONGUNS. co vs auc yoke hee tec Se el eg ORL. 41

00 LEDS TTT TSS Sy SO nn ea grate Se a ee 41

ee eae ee dn ye Le, Pe oe a oe 42

1 Based on the observations and studies of an expedition organized under the joint sup-

port of the New York Academy of Sciences and the Insular Government of Porto Rico.

Manuscript received by the Editor 1 February, 1915.

(i)

y) ANNALS NEW YORK ACADEMY OF SCIENCES

Page

Minor structures .....-....---- Spe ae oe Vee Seer 43

Enterolithic structares 2.25065 ¢265 Jeet te -5 2 3... 43

Double cross-bedding ...... 22. 225.003 gets ose no «'s = 2 43

Special relief features... . 2.2.2.0. 52020 c enc eee wes see oe a we — AT-

Playas 2.6 20 ao 8 5 dee w Se ateim We or te ets et AT

Promontories (28 2 so cs ois Oe saeco Meese 47

TeTTACES .ccccedecdacs dec voucetuseeu tl eee ne 2s eee 48

Cuestas: (ovis cee So eee Sc eee ode aes + 3 ee 49

Peneplain . 2... 5 css cas cae oon cee seis pies a ee 50

Hlaystack hills ... 2.500.205 cc eee nee eee eee 2 oe 51

Mineral’ re@SOUTCES ... 22. 6 acc cee eee we ew a en oes © oe ee ee 53

FOld «2 ok cc cet cence we ecw woe ee wees Sle ee ees eee 54

Copper ss ic ssa c ces cnc coe ete om cee ce = Uieceye Se at ie ee 55

Zine, lead and Silver... ....0.0ceeecececes ss => ee 55

ErOn os. wee ce dc os ee wo tne Slee eet 2 eee Be ee

Coal and Oil... sce ccs cen we awe eos ede Se 56

Tamerock .. 2.2. o000 ss 0eeensscdnceneeeeeee= =e 56

GUANO ...c ccc c cece ce wi cee eee es 6 ee i oe al eee tele 57

Road metal ....... 0.000 cce eens ccnns cee occe = ae ieee 57

Hot-Springs «oc i. See ss es ee ae ee eee PE 57.

Historical statement <2 2:2 “Sects lose = eens eee eer 2 ow 58

Future problems. ....-...:..i..-cee2-ceee oe descdnss es oes ee 61

Base. map: is. 0is te. oe Se eee eerer 61

Geologic MAP .........0e ce ewe cee e tee ccc es eene eee wee ee ee

District studies .. 063. cust cece ows ook eS bes banc orca 62

Reef-building organisms ......0.5.0. 0.00. c0s0ceu cee sm gee 63

Paleontology 220%... ss dane cde ae ee eee 2 oe Sacer 64

Tertiary subdivision. ... 2.5.0... 00.000 se ce ec oan os oe a eee 64

San Juan formation.» ... 2.502. 26 se < awn me auwer le ae ee 64

Subdivision of pre-Tertiary complex...........2...<.2.+s.00e—eeeene 64

Mineral reSOUFCES 2... oa ccc ec eek chee ae sem ood ee ane 2 eee 64

Petrography ... 2. seco eee we ls ed De ee Se ee eee 65

Physiography ......s000cteccest wen'es nce elec eae s cle Pres oF

Thermal. waters... 0.3% Soest Sc ee ee eae ss a ee oe 65

Geologic history .........0. 2.0. se cecc ecu ce se e- © ccs ieee eee

Collections ... 2.2.6... cso c Oeenwies cen eble Jecleecelenel SS 66

Tilustrations. ... 2... 5065. lec ad ce cen = demas be © aoe oe ee 67

Cross-sections . o.oo... jcc can cee cake ce cums enters one aeee an 67

Maps 2. 0. is nse Wubi ese otek ene ee je eee es oe 68

Acknowledgments <2 ss5552555208 deen I 68

Bibliography .... 220.60. cednn les Yee d oe a poe eee ee 69

INTRODUCTION

The Island of Porto Rico has never had a thorough or detailed geo-

logical study. There have been, however, a number of papers written

that have described special features or general conditions in a very ac-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 3

ceptable manner. Most of the writings? of this kind are widely scattered

in volumes of periodicals or proceedings of learned societies or pamphlets

which in most cases are not readily obtained or consulted.

From a perusal of these articles, one learns that Porto Rico belongs

structurally and genetically to the mountain chain now represented by

the isolated islands forming the principal West Indian group. Enough

work has been done, especially by R. T. Hill? to outline roughly the

geological history of the Island of Porto Rico and indicate on a map the

distribution of some of the formations. An especially good general de-

scription of physical features, also, is given by H. M. Wilson.* An intro-

ductory general description, in large part along the same lines as these,

will probably serve the present purpose.

GENERAL DESCRIPTION.

The Island of Porto Rico is situated in the Torrid Zone between lati-

tude 17° 54” and 18° 30” north and longitude 65° 13” and 67° 15” west.

It is the easternmost and southernmost of the Greater Antilles. It lies

within the trade-wind belt, and the constancy of these winds gives the

island a remarkably mild and uniform climate. There is an abundance

of rainfall on the windward side, which in this case is the east end and

the north side as far west as Camuy. The effect of the mountains across

which these winds blow is to make the south side of the island and most

of the western portion comparatively arid. Some districts are said to

have no rainfall for a whole year at a stretch.

The Atlantic Ocean lies to the north and east, the Caribbean Sea lies

to the south, while Mona Channel on the west separates the Island of

Porto Rico from Hayti. Brownson Deep, reaching the profound depth of

twenty-four thousand feet below sea level, one of the deepest spots known,

hes immediately to the north. Tanner Deep hes to the south, reaching

a depth of fifteen thousand feet. Although the relief of the island above

sea level is less than four thousand feet, this represents only the extreme

top of a great mountain mass which rises above the submerged platform,

from which its real height should be measured. The extreme relief differ-

ence represented by the summit of El Yunque on the one hand and Brown-

2The writings referred to, together with others that have some description of physical

conditions in Porto Rico, are listed at the end of this paper. Those of most usefulness

in the present investigation are certain papers by R. T. Hill and H. M. Wilson, besides a

very few others of less extended character.

3’R. T. Hitt: ‘Porto Rico.” National Geographic Magazine, volume 10, pages 93 to

fae (1889).

4H. M. Wiutson: “Water Resources of Porto Rico.” Water Supply Paper No. 39,

U. S. Geological Survey.

4. ANNALS NEW YORK ACADEMY OF SCIENCES

son Deep cn the other is approximately twenty-eight thousand feet. Re-

garded in this way, the Island of Porto Rico belongs to one of the higher

relief features of the earth.

It is in reality a badly eroded summit of a great mountain belonging

to an east-west chain or range including the Greater Antilles. The gen-

eral structural features of the islands are consistent with this east-west

axial trend which is expressed in the topography of the central Cordillera

extending from the west end near Rincon, where it starts abruptly from

the water’s edge, to Fajardo, where it terminates in E] Yunque, the high-

est point on the island. The mountain range, however, is not so simple as

this statement would lead one to believe, for there are in reality two

ranges or branches toward the east, one of which is known as the Sierra

de Luquillo, culminating in El Yunque, and the other, which is best de-

veloped in the divide between Cayey and Guayama, is called the Sierra de

Cayey. The military road crosses this latter branch near Aibonito over a

pass that reaches above 2,000 feet. ‘To the westward, the double character

of the mountain ranges is not so pronounced, but there is a semblance of

it in the spurs that reach the sea rather abruptly, one near Rincon and

the other near Mayaguez. The exact elevations of the higher mountains

have not been accurately determined, the values given on the older maps

being evidently too great. .The revised approximate elevations are: for

El Yunque, at the eastern end of the island, 3,750 feet; for El Guularte,

which stands to.the west of the Arecibo road, 3,610 feet. The highest

point in the Sierra de Cayey is about 3,000 feet. Many points are nearly

as high as those given, and all of the roads that cross the island reach

elevations on the divide that are in excess of 2,000 feet. Many of the

roadways in the interior districts reach elevations over 2,225 feet.

Although the island as a whole has a mountainous aspect, and although

much of the interior is very rugged and picturesque, there is usually a

comparatively gentle or smooth topography along the coast, and some of

the marginal areas are almost perfectly flat. These are uniformly at the

mouths of the larger rivers and represent river alluvium or delta-like

deposits,—they are known in the island as playas.

The aspect of the island as a whole is moderately rugged. Although

the major portion of the rock makeup is igneous, and although there is

considerable complexity of structure represented in all parts of the range,

all of the surface forms are of erosional origin. The relief is that of

early maturity in the interior and perhaps late maturity in portions of

the coastal districts. Exceedingly steep slopes are the rule in all parts

of the island where there is any considerable relief, and one of the most

surprising things is the way the soil clings to these slopes. One often

meee

BERKEY, GEOLOGICAL RECONNOISSANCE OF

SAN JUAN

ee,

RELIEF MAP

PORTO RICO

compiled by

PORTO

Role

3 drawn by

O.A.LJUNGSTEDT

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Via, Lee

CaB

RICO

__ 20miles

Porro Rico

I.—Rruibr Mar or

PLATE

)

Agriculture,

Department of

5 of the Forestry Division, l

)

Reproduced by permission from Bulletin ‘

6 ANNALS NEW YORK ACADEMY OF SCIENCES

sees slopes of this kind with angles of thirty to forty degrees from the

horizontal under cultivation.

The islands of Culebra and rasa he to the east of Porto Rico and

are said to be similar in structure and makeup, but neither of them was

visited on this expedition.

The streams of the island are numerous and surprisingly large for the

size of the areas drained. According to Mr. Wilson’s description® of the

water supply there are twelve large streams flowing north, four flowing

west, seventeen flowing south and six flowing east. Besides these there are

said to be 1,300 small streams, and on account of the heavy rainfall on

portions of the island many of them are of larger size than such an area

would usually afford. The main divide runs near the southerly margin

of the island, so that about one-third of the drainage is tributary to the

Caribbean Sea on the south, and about two-thirds to the Atlantic Ocean

on the north. This unsymmetrical position of the dividing range is an

abnormal feature, the cause of which is the subject of discussion in an-

other portion of the report. Because of the prevailing trade winds, the

rainfall is very unevenly distributed. The east end and the north side

are comparatively humid and plentifully watered; in contrast, the west

end and especially the south side are comparatively arid. In order to

overcome partly the shortage of water, a large system of irrigation 1s now

being developed on the south side of the island. At the east end, north-

east of El Yunque, there is an annual rainfall of 123 inches. On the

other hand, at Cabo Rojo, at the other extreme on the southwest side of

the island, it is exceedingly dry, and in occasional years there is said to

be not a single drop of rainfall. Other parts of the island vary between

these extremes. The wettest months are September and November.

In most districts, the underlying rock is compact enough to discourage

much deep water circulation and the stream run-off is correspondingly

responsive to the rainfall. In the northwest corner of the island, on the

broad limestone belt extending from Aguadilla to Camuy, there is a pre-

vailing tendency for the surface waters to sink into underground chan-

nels, leaving the surface very much more poorly watered than even the

somewhat scanty rainfall would lead one to expect. In some cases, streams

developed on the more compact rocks of the interior districts completely

lose themselves in underground channels upon entering the limestone

belt, and in some cases do not again come to the surface for several miles.

Elaborate caves and channel-like caverns are common and, in the north-

ern belt of limestone country, there are thousands of such occurrences

5H. M. WILson: “Porto Rico; its Topography and Aspects.” Jour. Am. Geog. Soc.,

Vol. 32; p. 220; 1900:

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO Y

still preserved that are now wholly abandoned by the waters that for-

merly occupied and helped to form them. There are no inland lakes, but

there are a few coastal lakes and they appear to be related to the develop-

ment of alluvial plains or playas and recent elevation and subsidence

changes.

The climate is strictly tropical, but it is so tempered by reason of the

constant trade wind breezes from the ocean and the elevated character of

much of the ground that it is usually agreeable and mild. The lack of

great changes of temperature and the prevailingly moist conditions on

most of the island have direct influence on the character of the rock decay

and disintegration and also on the quality of the soil produced as well as

its behavior as a residuary product. Other matters of climatic conditions

have little or no bearing on geological problems and may well be avoided.

The average daily temperature is eighty degrees; it rarely goes above

ninety degrees or below seventy. The maximum temperature is ninety-

nine degrees. Extremes recorded for the year indicate a range of forty

degrees.

The area of Porto Rico is given as 3,670 square miles, which is about

three-fourths the size of the State of Connecticut. It is roughly rectan-

_ gular in outlime and in actual dimensions is about thirty-five miles in

average width, and one hundred and five miles long from east to west.

It is the fourth in size of the West India Islands and is one of the most

productive and densely populated districts in America.

Because of the greater interest recently taken in studying the natural

~resources and natural history of Porto Rico, it was judged to be a suitable

time to make a more elaborate and detailed study of the island’s geo-

logical framework and history. In accord with this view the New York

Academy of Sciences organized an expedition which spent a part of the

summer of 1914 on the island. The accompanying descriptions are based

on the work accomplished by this expedition.

New York ACADEMY OF SCIENCES EXPEDITION

The geologists sent to make a preliminary study or reconnoissance of

the Island of Porto Rico left New York on the 15th of August, 1914.

Four weeks were spent in Porto Rico, the expedition returning to New

York City on the 21st of September. The party consisted of Dr. Charles

P. Berkey of Columbia University, New York, and Dr. Clarence N.

Fenner of the Geophysical Laboratory, Washington. Arrangements were

made with the bureau of transportation of the Insular government in

San Juan for conveyances, so that as much ground as possible could be

seen in the time available. More than 2,000 kilometers were covered by

ANNALS NEW YORK ACADEMY OF SCIENCES

the aid of this transportation service and observations were made in suffi-

cient detail to judge the general character and structural relations of the

formations crossed. In addition to this kind of travel on the roads, short

trips were made on foot to examine features or outcrops of rock which

appeared to deserve investigation, and an occasional more extended trip

on horseback was taken to points in the interior. With these facilities

for travel, it was possible for both members of the party to give undivided

attention to geological observations. It was possible to stop and make

Fic. 1.—Relief features characteristic of the interior ranges of Porto Rico

Photograph taken from the Ponce-Penuelas road at K-10, looking northward across

eroded formations of the older series to the main drainage divide.

brief examinations along all the roads at hundreds of places, and, on

several of the roads crossing the island, sufficiently elaborate data were

secured to furnish a basis for geological cross-sections showing both relief

and structural features. A complete circuit of the island was made and

in addition it was crossed from north to south on three principal roads.

This, together with numerous side trips into the interior, permitted ob-

servations to be made on practically every formation of any considerable

consequence in the island. No point in the whole area is situated more

than seven miles from some road or other point of observation covered

by the party, and, even in those cases, except in the extreme southwest

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 9

corner of the island, observations were made on all sides or completely

around the unexplored areas. Because of the extent of the reconnois-

sance, it is the opinion of.the writer that all of the essential, large, funda-

mental geological formational units have been found and their general

structural relations have been determined.

OBJECT OF THE EXPEDITION

The purpose of this expedition was, first of all, to determine the nature

and origin of the rock formations of Porto Rico, and to group them into

series suitable for use in subsequent geological work. In the second place,

it was the object of the party to determine as many of the larger struc-

tural relations as could be seen and to unravel as much of the geologic

history as such a hasty examination would warrant. In the third place,

the physiographic features were studied for the purpose of determining

their origin and relation to the formational structure and their bearing

on the more recent history of the island. Lastly, it was appreciated that

the island was complex enough to have many problems that could not be

solved without very much more extended investigation, and it was the

purpose of this expedition to point out the problems that should receive

special study and that seemed to give promise of important results. Con-

siderable attention has been given to the economic resources of the island

by private individuals and considerable money has been spent on various

enterprises connected with their development. These problems were also

kept in mind, and wherever convenient, special observations were made on

them. Although it is possible to make suggestions concerning these eco-

nomic resources, they are for the most part matters that should receive

very much more extended special study. A matter that concerns the wel-

fare of the island more directly than any of these is the question of

quality and variety and origin of the native soils. These of course are in

large part geologic matters also, and although this reconnoissance is not

sufficiently detailed to form the basis of a discussion of this matter, it is

one of the lines of investigation connected with further work that will

have direct value.

INVESTIGATIONS AND DISCUSSION

ROCK FORMATIONS

The most fundamental thing to be determined at the outset of an in-

vestigation of this kind is to discover and differentiate the different rock

types and the structural units to which they belong. All of the prelimi-

10 ANNALS NEW YORK ACADEMY OF SCIENCES

nary work of this expedition was devoted primarily to this question and

collections were made for comparison throughout the island. In the

beginning, observations were made chiefly along the coastal margins be-

cause of the greater amount of ground that could be covered and because

of the apparent simplicity of the outermost and younger or more recent

formations. The chief formations with their representative rock variety

will be discussed in order from the younger to the older series.

In the first place, such a reconnoissance shows that there are two great

series of formations separated by a marked unconformity. Both are

somewhat complex, but in that respect the older series is very much more

complex, both in range of composition and number of units involved and

in variety of structural relation, than the younger one. In spite of this

discrepancy, it is still the most convenient and useful division to make,

and, because of the strikingly different characteristics of the two series

and the great prominence of the structural break between them, there is

no possible chance for mistaking this fundamental feature.

The whole lot of formational units are therefore grouped under the

follow: , two heads:

1) Younger Series.

Including the Tertiary shales, reef limestones and recent deposits.

2) Older Series.

Including a complex group of formational units,—tuffs, ashes, shales,

conglomerates, limestones and a great variety of intrusives, all of which

are probably of pre-Tertiary age. 5

There are several possible subdivisions of the younger series, but in

this discussion only those exhibiting enough physical constancy and char-

acter to be useful in field correlation are taken into account. These are

especially (1) the San Juan Formation, a Pleistocene sand-dune deposit,

and (2) the Arecibo Formation, a series of reef limestones and associated

shales and marls. Besides these, there are more local developments that

deserve special discussion, such as the San Sebastian shales, the Juana

Diaz marls and sandy shales, and the Ponce chalk beds; but in a broad

grouping these are all phases of the larger Arecibo Formation and it will

take detailed paleontologic study to make the proper subdivisions.

The older series has many formational members and their general rela-

tions are reasonably well understood, but a systematic subdivision is not

yet attempted. Correlation in this series is still more difficult than in

the other, because of the great variation in character laterally and the

influence of igneous activities that prevailed throughout its whole devel-

opment. Some of the most characteristic of these types will be described.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO {jt

Younger Series

San Juan Formation.—On approaching the island by the usual route

entering San Juan harbor, the first rock whose structural detail can be

seen is that on which the city of San Juan itself is built. One can see

that the formation is made up of strongly bedded material that has all of

the structural characteristics of a cross-bedded sandstone and is resistant

enough so that it forms, at this point, a promontory extending far beyond

the supporting mainland and presenting a cliff face of at least 75 feet in

height. A closer inspection supports all of these observations as to

structure and adds the important observation that the granular material

Fic. 2.—Partially destroyed dune sand deposits of the San Juan formation

These deposits are a short distance west of Arecibo and are nearly covered with fresh

j dune sands of the same material.

is, in large part, calcareous in composition and organic in primary origin.

The same type of material, with all of its characteristic structures, was

seen at several other points along the north coast. Special studies were

also made on sea cliffs of this formation in the vicinity ‘of Arecibo, where

the exposures were so well developed that good photographs of the struc-

ture could be taken. One of these is reproduced as an illustration of the

characteristics of this formation. It shows exceedingly steep cross-bed-

ding structure that measures up to thirty-three degrees dip, and there are

also occasional structural lines that are almost horizontal. The rock is

exceedingly porous, the grains are unusually uniform in size, and the

12 ANNALS NEW YORK ACADEMY OF SCIENCES

binding material is calcareous, attaching one grain to another merely at

the point of contact. The extension of ledges of this kind of rock far

beyond the possible reach of swift-flowing streams, together with the fact

that the distribution is limited to certain sections of the north coast, and,

in addition, the evidence furnished by the internal structure of the rock

itself, lead to the conclusion that the formation is essentially an old dune-

sand deposit. Sand dunes are developed on the present coast line from

very similar material, but none of the very recent dunes are solidified.

On the other hand, the San Juan formation seems to have been developed

before certain of the later elevations and subsidences that affected the

island in its recent history, so that its material is fairly well cemented

and its base extends below the present water level. Its outcrops also ex-

tend to greater elevation above sea level than any of the modern dunes.

It is judged, therefore, that this particular formation is the most recent

of all in the island to act as a ledge former, and it is judged by its situa-

tion and content to be of Pleistocene age. See additional description

under heading “Structural Features.”

This is the most unusual formation in the island. It is a type seldom

seen or seldom recognized, and it is one of the smallest in Porto Rico, in

spite of the fact that it makes such an important showing at San Juan

harbor.

Because of the prominence of the formation in the city of San Juan it

is suggested by the writer that the name San Juan formation be used for

it and that this name be confined to the Pleistocene beds representing

solidified sand-dune deposits.

Arecibo Formation.—Next below and older than the San Juan forma-

tion is a great series of reef limestones and shell limestones preceded by

shales that form a belt of considerable width along the north coast and a

portion of the south coast of the island. In a large way this series forms

a structural unit. Above it in all cases le the recent alluvial -deposits

and the San Juan formation and below it le the older and more compl-

cated igneous and sedimentary rocks. The break between these two rep-

resents the chief unconformity in the whole geological column. The

heaviest development of this formation is along the north coast between

Tao Alto and Aguidilla. In this belt, the massive limestones of the Are-

cibo formation attain the greatest thickness observed anywhere on the

island, but no opportunity was found for determining the amount accu-

rately. There is in sight, however, certainly as much as 500 or 600 feet

in the bluffs along the Arecibo River. In this belt also, especially farther

toward the west, in the vicinity of Lares and San Sebastian, there are un-

derlying shales of considerable thickness which in places carry lignitie

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 413

material and which have been the object of some exploratory work for

coal. It is evident, from observations made, that the shale beds of the

Arecibo formation vary greatly and in some places are entirely wanting.

At the Arecibo River, for example, where the beds of the formation can

be seen well exposed in the river bluffs, there is no shale development at

all. ‘The limestones he abruptly on the eroded and somewhat weathered

surface of the older formations which at this point are represented by

coarse and obscurely bedded or even massive volcanic tuffs. At the best

place seen, which was on the east side of the river about opposite K-66 on

the Arecibo road, there were a few feet of transition material between the

limestone beds proper and the unmodified tuff. It had not the structural

appearance and makeup of the shale as seen at other points, however,

and this part of the formation is regarded as entirely absent on the Are-

_¢cibo River. Shaly beds, however, are seen again on the south side of the

island and their best development is in the vicinity of Juana Diaz, where

some of the beds are distinctly sandy and rather fossiliferous and carry

petrified wood. Lignite is also reported from this vicinity, but no ma-

terial of that kind was found by the writer. It is more than likely. that

the shale beds on opposite sides of the island do not correspond in horizon

at all, but that structurally they are both basal beds.

The most striking development of the shale beds and overlying marls

and softer layers of thin-bedded character, instead of the massive reef

structure, is on the Jacaguas River south of Juana Diaz. The dips also

of the formation in this particular locality are much greater than those

observed at any other point. For considerable distances an average dip

of 30 to 36 degrees was estimated and the total thickness represented,

based upon the width of the belt, must be at least 3,000 feet. At no

other point on the south side, however, was there an opportunity to see

whether the beds of this character are constant or of large lateral extent.

As one goes eastward, a comparatively short distance, they are almost en-

tirely lacking. On the Descalabrado River, which is only ten kilometers

to the east, the underlying older series of tuffs and intruded shales and

limestones were followed to a distance of two miles south of the military

road, whereas at Juana Diaz the basal shales of the Arecibo formation

begin a half mile above the military road. It appears, therefore, that the

formational margin is swinging rapidly southward, and it is judged,

from other observations made, that there is almost nothing of it repre-

sented at a distance of twenty kilometers to the east, or, in other words,

that the formation does not extend farther east than the vicinity of Sa-

linas. At one other point on the south margin of the island, there is an

unusually good opportunity to follow the successions of formations, and

14 ANNALS NEW YORK ACADEMY OF SCIENCES

that is in the vicinity of Guanica. Limestones belonging to the older

series occur immediately south of Yauco and are very strongly developed

there. The hills in which these beds outcrop extend southward almost

continuously to Guanica, but at that point observations showed that the

formation had changed and is actually part of the reef limestone of the

Arecibo formation, although it is possible that the large fault observed

west of Ponce may pass through this area and obscure the other structural

relations. It looks, from the rapid survey, as though it would be favor-

able for some of these additional studies of the character of the lower

beds of the Arecibo formation. Between Guanica and Juana Diaz, wher-

ever the inner margin of the Arecibo formation was seen, it was bounded

by a fault which brings the upper beds abruptly against the older tufts

and shales of the pre-Tertiary.

The formation furnishes an abundance of fossils. The lower portion |

or the lower beds on the south side of the island, as seen at Juana Diaz,

seem to be the most promising for a determination of the age of the beds

of the. formation. Higher beds, forming a chalky white limestone to the

west of Ponce, are also very fossiliferous, but in this area they are sepa-

rated from the older rock series by a fault, so that it is quite impossible

to tell how far above the base of the formation these beds may lie. It is

judged that the portion of the formation seen at Guanica is a still higher

horizon, but the exact age values have not been worked out. The total

thickness of the whole Tertiary series on the south side of the island is

very great. It was estimated that the shales and marls and limestones

in the vicinity of Juana Diaz must certainly amount to three or four

thousand feet. For a long distance along the Jacaguas River south of

Juana Diaz, the beds stand with a dip of approximately thirty-five de-

erees toward the south and throughout the greater portion the character

is notably different from the beds occurring farther to the west which are

judged to overly them. |

To the east of San Juan, along the north coast, there is much less

prominence of the Arecibo formation and after passing Rio Piedras it in

no place crosses the main road. ‘There are occasional hills somewhat

similar to those characteristic of the landscape of Bayamon and vicinity,

but they do not reach to so great a height and are separated by very much

larger stretches of low ground. The strongest development of this for-

mation seen to the east of San Juan is that along the Grande de Loiza

River between the railway and the coast. In going still farther east to

the vicinity of Luquillo, the inner margin of the formation passes out to

sea and the older formation reaches the shore. From this point around

the whole eastern end of the island no more outcrops of the Arecibo for-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO. 15

mation were seen, and this is true also of the southerly side coming from

the east end through Naguabo and Yabucoa and Guayama, and still far-

ther west to some undetermined point between Guayama and Santa Isa-

bel. There may he, however, occurrences of this formation underlying

the alluvial material along the coast at points considerably nearer Guay-

ama than the outcrop map indicates.

The formation as a whole is essentially a structural unit. Although

it is quite easy to distinguish the underlying shale member, especially

well developed between San Sebastian and Lares, and although there are

other structural changes, there is nowhere any appearance of unconform-

ity or marked break in the succession. Besides, the shale member is not

everywhere developed and, as a matter of fact, is seldom seen in tracing

the formational boundaries. In some places, it is definitely shown to be

absent, so that it seems unwarranted to represent this member as an im-

portant part of the formation, so far as areal distribution is concerned.

Judging from field observations already made, the shale and mar] beds

are more extensively developed on the south side than elsewhere. But

this is based on observations in one particular area and the member can-

not be traced very far in either direction because of other difficulties.

The beds lying above the shales and representing the part referred to as

a more massive limestone portion are probably susceptible to considerable

differentiation on the basis of fossil content, and it is entirely possible

that a rather complete range of Tertiary horizons may be determined

after complete paleontologic study. In the field, however, and on the

basis of structural factors, there is no apparent ground for subdivision.

In this discussion, therefore, the whole series of beds, from the uncon-

formity at the base to the alluvium and San Juan formation overlying it,

is referred to as a single formation and, on account of its extensive devel-

opment in the region about Arecibo, it is suggested that a suitable name

would be the Arecibo formation.

_ Some parts of the formation show the peculiarities and content of a

coral reef, and these portions have the most irregular and most massive

structures. Other parts show bedding structure more or less perfectly de-

veloped, and throughout the whole formation here and there, at irregular

intervals, and usually of only very limited extent, there are more shaly

facies. It is the opinion of the writer that this irregular distribution of

shaly beds is responsible for one of the peculiar topographic features de-

veloped in the belt underlain by the Arecibo formation. This is the

occurrence of almost perfectly flat soil-covered areas of no very great

lateral extent at different levels, above which the numerous small knobs

or hillocks of limestone rise, giving the peculiar haystack-like topography.

1G. ° ANNALS NEW YORK ACADEMY OF SCIENCES

This type of topography is represented by the small level tract surrounded

or dotted over with small hills, called “pepino hills” locally, standing

like haystacks above the plain at many different elevations above the sea.

This leads to the belef that the fundamental control in its development

is the presence of a shaly bed mm the series, which forms at each poimt the

basis of the local plain. The hillocks standing above it or surrounding

it represent remnants of the more massive and probably more porous and

more easily destroyed limestone which has been attacked and largely

removed by weathering, and especially by solution, down to the more

" £

ites: SY ta es

Fic. 3.—Unconformity telow the Arecibo fermatian

This view shows the older tilted and eroded tuffs and shales below, as seen on the

Arecibo River. The contact is immediately beneath the horizontally bedded limestones at

about the center of the view where the chief weathering is noted.

resistant shaly soil-forming member. If this is the principal cause of

the peculiar topographic form just described, it is quite easy to see that

the distribution of such features should be expected to be rather irregular

both in Jateral extent and in actual elevation above sea level or in rela-

tion to the different horizons in the formation itself. In regarding this

as the principal factor, there is no tendency to overlook the fact that the

island has stood at different elevations with respect to sea level in former

times, and that a corresponding difference In ground water levels would

be felt throughout the border region. But there is no evidence whatever

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 17%

‘that the numerous different levels represented by many of the small

plains referred to above were necessarily connected with any of these

subsidence changes. Additional comment on the hillock topography, so

strongly developed in some parts of the area, is made in connection with

discussion of drainage in another part of this paper.

The Arecibo formation is of Tertiary age. So far as identifications of

the fossils have gone, they appear to confirm the opinion that the larger

part of the formation belongs to the Oligocene epoch. These determina-

tions were based largely on collections made in the heavy limestone beds

and reefs in the vicinity of the Quelbradillas River. The shale beds

lying at the base of the series, and exposed farther to the south in the

vicinity of Lares, are certainly somewhat older and probably belong to

the Eocene epoch. There are higher beds developed rather irregularly

that doubtless represent still later time, referred by Hill to the Miocene

epoch, but these determinations must be left to future detailed study of

the formation as a whole.

It is considered eminently fitting to refer to some of the chief varia-

tions which have especially strong development in certain localities, by

special locality designations, such as,—San Sebastian shales, Ponce chalky

limestones and marls, Juana Diaz shales and marls, Guanica coral reefs,

Quebradillas reef Hmestones, etc.

The correlating of all these and other local representatives of the Are-

cibo formation is a work that can be done only by extensive and detailed

stratigraphic study and paleontologic comparison. This is one of the

larger pure-science problems awaiting future investigation.

Older Series

Below the Arecibo formation and forming the surface in the interior,

beyond the Arecibo margin, the island is made up of an exceedingly com-

plex series of many different kinds of rocks. They include chiefly varie-

ties of igneous rocks, both extrusive and intrusive, both fragmental and

massive, ranging from small stringers or dikes or flows to large boss-like

masses that cover many square miles in area. In addition to the igneous

rocks of these types, there are numerous shale beds and conglomerates of

rather massive habit aggregating a very great thickness, and with them

are associated limestones and foraminiferal beds of considerable variety.

A study of the rocks of this series for the purpose of determining their

character and origin indicates that practically everything in the older

series except the limy portions of the shales or the limestones proper are

more or less directly of igneous origin. The coarser materials and those

least affected by any secondary processes are the tuffs which are of direct

18 ANNALS NEW YORK ACADEMY OF. SCIENCES

voleanic origin and are exceedingly abundant and extensive. They are

found at intervals in all parts of the series, and it is impossible to say

that they are either more or less abundant in those portions which appear

to be older, rather than in those which appear to be younger or higher in

the series. ‘The closest associate of these materials is the bedded tuff,

made up of volcanic fragments which have been somewhat assorted by

surface agencies so that they exhibit some sedimentary structural charac-

teristics. These are also exceedingly abundant and widely distributed and

they pass by insensible gradations of finer and finer materials into those

Fic. 4.—Typical shale occurrence

This is seen along the road between Ponce and Penuelas at K-10. The beds at this point

lie in a less disturbed attitude than is usual in members of the older series.

that are recognized as true ash beds. Most of these have become so thor-

oughly cemented, or so much modified by secondary attack, that they now

present a perfectly sound and compact appearance. In thin section,

however, it is easy to see that the material is wholly voleanie and that

the bedding is the only secondary modification except that having to do

with the binding, induration or alteration of the rock. The ash beds are

probably close relatives of the so-called shales.

Shales——Rocks of this type are developed characteristically at Fa-

jardo, at Mayaguez near Baranquitas and at numerous other points, espe-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO {19

cially along the divides toward the west. At the two points first men-

tioned, in particular, they are light yellowish or reddish in color, rather

porous in structure, strongly bedded and have a prominent blocky frac-

ture habit. The exact character of these rocks is a question under study

at the present time, but enough has been done to show that, in their

present condition, they have been oxidized to the yellow or red color, and

have been leached so that they have a porous structure and hght weight

due to the removal of at least a part of some constituent that is more

readily soluble than the rest of the rock. Microscopic comparison with

beds of the same structural relation, but of very dark color and very

dense habit, leads to the opinion that the two types are not essentially

different in origin, but that the lighter colored and lighter weight shales,

such as are found at Fajardo and at Mayaguez, are simply the weathered

equivalents of darker ones. It appears from this comparison that the

shales are normally highly calcareous and that the lime content is sup-

plied by the presence of a very large amount of organic matter in the

form of foraminifera. In some cases this organic matter makes up fully

one half of the rock and in all cases weathering produces a very porous

effect that should be expected to be identical with the red and yellow

shales occurring typically at Fajardo and Mayaguez. The siliceous con-

tent of all of the shales examined proves to be exceedingly fine and wholly

lacking in granular or quartzose material such as characterizes most sedi-

mentary shales. It is the judgment of the writer that this material in

the shales of Porto Rico, instead of being the ordinary disintegration

products derived from the weathering of ordinary land masses, is in

reality largely ashy material of volcanic origin. With this conception of

them, it would appear that even the limy shales are therefore close rela-

tives of the ash beds, and it is entirely possible that they do not represent

any great difference in history, but rather somewhat different surround-

ings during accumulation.

. Limestones.—Besides the shales, there are massive limestone beds of

several different types.

In most cases the occurrences are separated by structural complexities

that make it uncertain about field correlation, but undoubtedly later field

study will connect some of these and additional paleontologic study will

arrange their succession. The most prominent occurrences seen are de-

‘seribed below.

Ooamo Tuff-Limestone-—The limestone with the closest genetic re-

semblance to the types already described is represented in a broad belt

passing from south to northwest across the upper end of Coamo Reservoir

~ near Coamo Springs, and which can be traced in prominent development

20 ANNALS NEW YORK ACADEMY OF SCIENCES

westward across the Descalabrado River to the Jacaguas Reservoir. Sim-

ilar limestones are found at other points on the south side of the island

and are judged to belong to the same member of the older series. Be-

cause of the strong development in the vicinity of Coamo Springs, and

because of the fact that it represents a type so striking as to be recog-

nizable as a field unit, the limestone has been called by us in the field the

Coamo Tuff-Limestone. It is developed in the vicinity of the Coamo

Reservoir to a thickness of several hundred feet and its most character-

istic appearance is the brownish mottled color effect produced by the

Fic. 5.—IJnterbedded coamo limestone layers with massive tuffs

This formation is seen at the military road crossing of the Descalabrado River.

presence of fragments of tuff and accumulations of ash. In some beds

this material is so abundant as to make up almost the whole rock and it

becomes an interbedded tuff layer. Occasionally the limestone beds are

very pure and almost entirely free from volcanic materials, and there are

also numerous beds of real volcanic tuff, but typically there is an inter-

mixture of tuff materials with the lime in great enough abundance to

give a brownish spotted or mottled effect. An equally characteristic

feature of the rock is its concretionary or nodular appearance due to

algous growths to which the lime accumulation is chiefly due. The finest

development to be seen anywhere in the island is on the Descalabrado

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 2{

River at the point where the military road crosses it. A photograph of the

interbedded relation of tuff and limestone at this point accompanies this

discussion. This is probably, in part at least, the rock referred to by

Hill in some of his discussions as “mountain limestone.” It seems to us

that the several occurrences of limestone beds which have clearly differ-

ent relations in the series, and the probability of being able to place them

in different horizons, based on this content, makes it desirable to use

more characteristic local designations for them. Such usage cannot be

confusing even if it should be proven, as it may very well be, that some

of them are identical. In this discussion, therefore, wherever possible,

the chief occurrences that are not clearly identical with formations

already described are characterized by adding the name of the locality

-where there is especially good development of the formation. <A lime-

stone seen at Coqui, considerably farther east, and several occurrences

near Yanco, and others still farther toward the west, have many points

of resemblance to the Coamo limestone. In some cases even the brown.

volcanic spots are also present, but this is not true of all places. The

most constant index as seen in the field is a fine meshed coralline fossil

form, strikingly resembling a piece of loosely woven cloth.

The limestone beds developed at Coamo reservoir, however, rarely show

this type, but instead have a remarkable development of alge of the form

known as Lithothamnia.

Trujulo Alto Limestone.—There are several other limestone members

in the older series. One has been observed only on the north side of the

island in the vicinity north of Trujillo Alto and in the vicinity of Loiza.

' This is a very dense fine bluish limestone made up wholly of fine micro-

scopic organic growths. In some places it has a rough fragmental struc-

ture, but for the most part the rock is massive and the abundant organic

content, largely alge, is its most striking characteristic. Whether it is

younger or older than the Coamo limestone has not been determined, but

that it belongs essentially to the same general series is quite certain.

On account of its distribution it is conveniently referred to as the T’ru-

jullo Alto limestone recf. This member probably has a very moderate

thickness and no great areal-distribution. It is affected by solution de-

veloping caves at the Trujillo Alto locality.in much the same manner as

is the Arecibo formation, but this rock is a much more compact type and

its content and structural relations are quite distinct. It was probably

of reef origin also, but is associated intimately with the upper shale

members of the older series rather than with the Tertiary series.

“Shred” Limestone.—Another limestone may be seen at several places

on the Arecibo-Ponce road on the south side of the divide from K-13 to

29 ANNALS NEW YORK ACADEMY OF SCIENCES

K-19. It has no great thickness, but there are several independent beds.

Its most striking character is the presence of patches of dark color dis-

tributed in a shred-like way through the grayish mass. The rock as a

whole is massive, exceedingly compact, of a bluish gray color, and, except

for these dark-colored shreds, shows no recognizable structure whatever.

They contain alge, however, which are expected to determine something

more definite about their position in the series, but they are obscure

forms, and doubtless considerable work will have to be put on these beds

to determine their exact horizon. They are intimately associated with a

series of igneous fragmental beds and a considerable thickness of very red

shale or ash beds. ‘Together these alternating limestone and red frag-

mental beds make a striking structural succession which was not seen

anywhere else. They lie in a position which is not far from the point

where the Coamo limestone belt should be expected to cross the Ponce-

Arecibo road, but no such structural development has been noted at any

other place. It is, of course, possible that the fine red ash represents the

tuffs of the regular Coamo formation and the “shred” hmestone repre-

sents a phase of the Coamo not developed elsewhere, but the striking

physical difference encourages the making of a distinction, at least for the

present.

La Muda Ivmestone.—A rather heavy development of limestone in the

vicinity of La Muda between Rio Piedras and Caguas has some super-

ficial resemblance to the beds just described from the Arecibo road, but

their relationship is not fully determined. ‘The rock is not prominently

tufaceous and is not marked in the same way. It has in places a coarse

fragmental structure almost completely obscured by healing and it is, as

usual, attacked by cave development. Some of the caves have collapsed,

leaving a complex aggregate partly made up of igneous material filling

the former chambers. A conglomerate bed lies below the limestone and

shale at this point and both are cut off abruptly by a large intrusive

mass. How these are related to other typical members of the older series

is not known, but it will be possible to trace the beds to more definite

relations. This is probably one of the oldest limestone members in the

pre-Tertiary series. It is conveniently referred to as the La Muda

Timestone.

In addition to this there are very numerous small or thin local devel-

opments of limestone layers distributed through the shale beds at various

points. These are taken to be, in most cases, simply somewhat more

heavily developed limy lavers of the same origin as the rest of the fora-

miniferal and ashy shales; but the nature of their origin shows that it is

reasonable to expect a development of calcareous content sufficient to

make them more of a limestone than a shale.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 23

“Mountain Limestone.”—In some localities, such as that near Barran-

quitas, and probably at other points along the same divide, the shaly beds

become very calcareous, as has been pointed out by Mr. Hill. These

were referred to by him as limestones and are probably included in his

“mountain limestone” and estimated that the thickness of such beds

amounted to fully a thousand feet. The prominence of the shaly struc-

ture in these beds and their apparently close genetic relation to the type

described in this discussion as shales, leads us to regard this occurrence

as essentially the calcareous extreme of the shale series. As already

noted, the shales are characteristically calcareous and some of them are

predominantly so.

Corozal Limestone.—At Corozal, a fragmental limestone was seen just

south of the village. Its relations were not worked out and its meaning

is therefore not well understood, but its structural peculiarities lead to

the suspicion that it may be associated with volcanic fragmental accumu-

lation. It was found that in at least one place in another part of the

island there had been volcanic outbreaks through heavy limestone beds,

and it is evident that fragmental material from such activity might

therefore include a good deal of simple limestone fragments. It is hardly

conceivable that they would in most places accumulate in enough abun-

dance to make a limestone bed a second time, but such a thing is doubt-

less possible. ;

Conglomerates.—There is a very extensive development of conglomer-

ate occurring in a belt whose general trend seems to be from southeast

toward the northwest, crossing the military road between Aibonito and

Coamo. There must be a total thickness of strata, including shales and

interbedded tuffs with occasional small lmestone, of perhaps several

thousand feet. In all parts of the formation where conglomerate is de-

veloped, the pebbles represent the same kinds of rocks as were encoun-

tered in the tuffs and intrusive masses. Actual representatives of previ-

ously solidified bedded material or indurated ash or shales are very rare,

but in one case at least a pebble was observed that was judged to repre-

sent a fragment from an older silicified tuff. As a matter of fact, the

materials are practically all of simple igneous character and the matrix

in most parts of the formation is very abundant, or even predominant,

the particles of which are of the same igneous material. The distribu-

tion of material and the range of composition leads one to believe that

this conglomerate represents a special state or condition whereby ma-

terials of essentially tufaceous origin were, immediately after their vol-

canic eruption, worn, rounded, somewhat assorted and bedded and mixed

with related material. At the point examined, there was no satisfactory

24 ANNALS NEW YORK ACADEMY OF SCIENCES

evidence of marked unconformity between the conglomerate above and

the underlying series of formations. But the fact that the conglomerate

beds, which follow to great thickness, are prevailingly of simpler struc-

tural habit, as compared with the calcareous shales, ash and tuff series

immediately below, suggests that there may be a break here of larger

consequence than is observed in other parts of the pre-Tertiary or older

series.

The development of so extensive a series of conglomerates doubtless

does represent a considerable change in physical conditions, compared to -

those controlling simpler deposits which preceded and followed them, and

it is possible that it may be found useful in separating the complex series

of older mixed bedded rocks and tuffs into an older and a younger divis-

ion by using this conglomerate as a dividing member. This is supported

to some extent by the occurrence of a conglomerate of similar character

but of very much less extent on the north side of the island, several miles

south of Bayamon, and also one of apparently less prominence near La

Muda. Ti additional field work should show that the conglomerate belt

could be traced from one side of the island to the other, it seems to me

that it would be entirely practicable to make this division.

The conglomerate is invaded by igneous intrusive material in much the

same manner as is observed in the other rock formations, but the massive

habit of the rock as a whole leads to a predominance of transverse dike-

like masses rather than the simpler looking sills. At one poimt in par-

ticular, however, near K-86 on the military road between Aibonito and

Coamo, the conglomerate has been invaded by a magma that must have

been fluid enough to penetrate the porous matrix surrounding the con-

glomerate pebbles where it now exhibits a crystalline habit. This in-

jected matrix is essentially a coarse diorite porphyry in composition,

through which the pebbles are distributed in the manner that they seem

to have had in the original rock, so that there are still obscure traces of

bedding structure. There are additional petrographic peculiarities in

this rock that will be described under a different heading. This tendency

of the dioritic magma to penetrate and incorporate fragmental matters -

was noted in several other places. It was most strikingly exhibited in

certain intrusive members cutting through tuffs and shales. In some of

these cases there is so great a quantity of fragmental matter as to wholly

obscure the true nature of the rock unless one can see the structural rela-

tions. In the case of the conglomerate, however, the crystalline habit of

the matrix is a striking feature and it is very evident that it is wholly

different from the regular conglomerate habit.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 25

Tuffs—The most abundant of all of the rock types is represented by a

great variety of volcanic tuffs. These accumulations seem to be the funda-

mental basis of the whole island. Whatever has been formed in the way

of shales, sandy beds or conglomerates or any other of the ordinary sedi-

mentary types seem to be directly derived from the same material that

makes up the tuffs and the rocks directly associated with them, related in

an interbedded succession. In most cases, the tuffs are essentially massive

in their structural habit and are made up of a complex accumulation of

large and small fragments of volcanic materials which prove to be chiefly

andesitic and closely related porphyritic types. There is great variety in

texture and minor structure and present condition, but on the whole there

is enough uniformity of actual composition to justify classing them all

as andesitic tufts. ,

In very many places there is obscure bedding structure indicating a

tendency to assort and work over this material at the time of its deposi-

tion. This is especially noticeable in the finer materials and some of

these beds are made up essentially of ash. These ash beds resemble the

type referred to as shales so closely, in some places, that it is impossible

to distinguish between them in their field appearance. I judge also that

there is practically a gradation from one rock to the other, the ash beds

showing transitions to shales, especially where they have developed under

conditions encouraging much weathering and working over of the frag-

mental materials and promoting the growth of organisms in sufficient

amount to make the accumulating beds somewhat calcareous in compo-

sition.

Tuffs and ashes are well known to be especially lable to attack by

alteration and to the ordinary changes that modify rocks. It so happens,

therefore, that many of these representatives are completely modified and

have become so dense that they exhibit none of their ashy or fragmental

structure without microscopic examination. In this condition they are

usually also exceedingly hard and as resistent to destruction as the hard-

est crystalline rock. The largest development of massive almost struc-

-tureless tuffs which were seen occupy the Sierra de Cayey between Guay-

ama and Cayey and also the range along the military road toward

Aibonito; but there are extensive occurrences in many other sections.

Some of the most prominently developed bedded tuffs and ash beds were

seen on the north flank of El Yunque along the Sabana River and on the

north side of the divide below Comerio, and also along the Ponce-Arecibo

Road both near the summit of the range and farther to the north midway

between Arecibo and Utuado.

From what was seen of this type of rock, it was not possible to form a

26 ANNALS NEW YORK ACADEMY OF SCIENCES

definite conclusion concerning the age represented except by their rela-

tion to certain interbedded shales and limestones. It appears that the

underlying older portions of the series of tuffs and ash beds have com-

paratively little of such interbedded calcareous material and have every-

where, been modified or altered or metamorphosed to a greater degree than

beds that lie higher in the series. But beyond this there is little to judge

of the actual age. As one goes higher in the series, however, there are

occasional prominent limestone members with which tuffs are intimately

associated or interbedded, and it may be possible, by reason of these rela-

tions, to form a more accurate estimate of the geologic age of this later

portion of the series.

Volcanic Flows.—In addition to the sedimentary beds of various sorts

and the related tuffs, there are at occasional places evidences of volcanic

lava flows. These were seen at several places on the road between Baya-

mon and Barranquitas. They are amygdaloidal in present habit and

represent vesicular basalt and andesites. On the whole, evidences of lava

flows on a large scale are wanting. This kind of product seems to have

been very much more rare than the fragmental type. A more prominent

thing as a structural feature is the occurrence of very numerous intrusive

bodies.

Intrusives.—The intrusive masses in Porto Rico occur in all parts of

the island and in all of the formations except the Arecibo and the over-

lying alluvial deposits. No such evidence was seen in any part of the

Tertiary of younger series; but the complex series of rocks representing

the pre-Tertiary, here referred to as the older series, are cut in all sorts

of ways by both large and small intrusive masses. The smaller intrusives

are chiefly andesite porphyry in composition and have everywhere pene-

trated the shales and ash beds. The commonest occurrence is in the form

of small sills or sheets conformable to the bedding structure and varymg

in thickness from only a few inches to many feet. These sills are so per-

fect in form, have so little disturbed or modified the adjacent beds, and

are so similar in general composition and appearance, after weathering,

to the associated sedimentary beds, that it is quite impossible to deter-

mine in all cases how much intrusive and how much original sedimentary

rock is involved. The only thing noticed as a rule is the uniformity of

petrographic structure that seems to be characteristic of the intrusive as

compared with the associated beds. The simplest occurrences of sills of

this kind, which at the same time show their igneous intrusive character,

were seen near Fajardo, near Rio Piedras and in the vicinity of Comerio.

But occurrences of the same kind are exceedingly numerous in nearly

every district and in total amount form a very great additional thickness

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 27

to the bedded rocks, shales, ash beds, ete., with which they are associated.

In some cases, these invading magmas have incorporated great quantities

of fragmental matter, giving the rock in its present condition a very

strikingly fragmental appearance. This habit associated with its per-

fectly apparent intrusive relation makes a very unusual combination in

the field. In many places there are included blocks of immediately adja-

cent rock such as one sees in the occurrence at the quarry at Fajardo

Playa, but, in extreme cases, the mass is chiefly fragmental in its make-

up and one could not readily interpret its history without complete field

Fic. 6.—Diorite porphyry sills

These sills are intruded between layers of calcareous shales and ash beds on the road

near Comerio. The streaked or banded layers are shales, the massive portions, seen best

at the left side of the print, are sills. A transgressive relation can be seen between the

two layers at the extreme left.

determination of its relations. Such occurrences may be seen in the

vicinity of Guayama on the road about a mile east of that place, and also

a short distance south of Rio Piedras.

Few of these intrusive masses show anything but a rather uniform

medium grain texture and larger ones have the average appearance of a

rather fine or medium grain diorite. The granular appearance, however,

is probably deceptive, due to the way the rock disintegrates, for thin sec-

tions made from many typical intrusive representatives are nearly all

plainly porphyritic in texture.

28 ANNALS NEW YORK ACADEMY OF SCIENCES

In addition to these intrusive masses, which are of small size or at least

of not very great areal dimensions, there are in a few districts large boss-

like occurrences of massive coarse-grained igneous rock. The boundaries

of none of these have been traced out. but it is certain from the distribu-

tion now known that in each case the area occupied is several square

miles in extent. The most prominent one of this type of intrusive mass

is that seen in the southeast portion of the island, including the district

about Huamacao and Yabucoa and Las Piedras and Juncos. . Whether or

not this is all one mass belonging to a single intrusion has not been deter-

mined. The variety of composition seen in the different samples taken

at different points is consistent with the presence of more than one intru-

sive unit; but it is also possible and quite as likely that the variety ob-

served is wholly due to magmatic differentiation. The southerly portion

of this mass, especially that near Yabucoa, is represented by a very coarse,

very guartzose and almost pegmatitic granite. Farther to the north, in

the vicinity of Las Piedras and Juncos, the rock has the appearance of a

syenite. Although a part of the rock does show the composition of a true

syenite, by far the greater number of specimens collected on this expedi-

tion show the presence of quartz in sufficient amount to make the rock a

granite. It would appear, therefore, that this occurrence in the south-

east portion of the island is essentially a granite mass and that it is of

unusually large size, reaching practically from the coast at Maunabo to

Caguas. The distance across this mass is, therefore, not less than about

12 miles north and south. In all probability it is not of quite so great an

extent east and west, but these boundaries are unknown.

One other large intrusive mass was observed in the west central part of

the island, in the vicinity of Jayuya and Utuado. Im general appearance

and texture this rock, in the average outcrop, does not differ much from

that seen at the east end of the island which is commonly referred to as

syenite. In this occurrence, however, such specimens as have been exam-

ined with the aid of the microscope, show the presence of quartz in most.

cases In sufficient amount to make the rock of granite composition.- In

this case, as in that referred above, there are considerable differences of

composition shown by the rocks which seem to be a part of the same mass.

Specimens found, for example, near Adjuntas have the compositional

characteristics of diorite, whereas a specimen taken near the margin of

the boss on its northerly side, near K-53 on the road toward Arecibo, is

a syenite. At certain other points near-Utuado, the rock is a granite

porphyry.

The best idea of the variety of composition and textural quality repre-

sented by all kinds of intrusives in the island can be gathered from an

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 99

examination of the water-washed pebbles in the stream beds that have

come down the steeper mountain sides. An examination of such material

shows an extremely large varietal range, and, although by all means the

greater number are some variety of diorite or andesite porphyry, there

are occasional more basaltic and more acid types represented.

Summary.—From this description of the variety of rock types repre-

sented by the leading field units, it may be readily seen that a subdivision

into members of mapable constancy is no easy matter.

It is the opinion of the writer that, for the present work and for in-

vestigations immediately to follow, local designations or names will be

of most direct usefulness, and that a correlation should be expected to be

the final outcome of a series of such studies. For the present, therefore,

it is judged best to use the terms Fajardo shales, Mayaguez shales, Jayuya

road shales, Barranquitas shales, etc., as suitable names in these respective

districts for the shale member of the older series, without any intention

of suggesting their equivalence. In spite of the physical similarity in

these cases, it is not at all likely that they belong to the same horizon.

Similar argument will hold for most of the other members,—the ash

beds, the tuffs, the limestones and the conglomerates,—and it is recom-

mended in these cases, also, that locality designations be used in the field

investigations. Those described in this report are not necessarily all that

deserve special designation, but the same rule may be applied to addi-

tional occurrences without in any way obscuring the ultimate solutions

of the problem of correlation. On account, therefore, of the complexity

of the structure and the limited amount of work vet done in connecting

the separated occurrences into continuous field units, it is advisable to

retain such terms as Coqui limestone, Coamo limestone, Corozal lime-

stone, La Muda limestone, Trujillo Alto hmestone, Cayey tuffs, etc., for

the earlier descriptions and special studies.

PETROGRAPHIC RANGE

There is an exceedingly great variety of certain classes of rocks in the

Island of Porto Rico. Those most prominently developed and showing

the greatest variation in minor character, structure and relationship are

the volcanics, especially the volcanic fragmentals. All sorts of tuffs, cin-

der beds, ashes, mud flows and bombs are represented in great quantity,

in very wide distribution and in all stages of alteration and induration.

Observations made on thousands of occurrences of this character of

materials leads to the conclusion that most of it is essentially of andesitic

composition. Although there is an occasional fragment of either more

basic or more acid composition, the predominant types are always of

30 ANNALS NEW YORK ACADEMY OF SCIENCES

andesitic makeup. ‘The present condition of these rocks, representing as

they do nearly all stages between fresh material and either a thoroughly

weathered or considerably metamorphosed condition, is a more interest-

ing study than their primary composition. Some of the most dense and

resistant rocks in the whole island are these older metamorphosed tuffs

and ashes, :

Next in point of abundance is the group of crystalline igneous rocks.

In this case there is somewhat greater prominence of varieties represent-

Fic. 7.—Photomicrograph of a typical thoroughly indurated andesitic ash, magnification

28 diameters

A rock of this type appears in the field as a dark-greenish hard resistant obscurely

bedded layer, usually closely associated with more massive tuffs on the one hand or

more strongly bedded shales on the other. The clear grains are mineral fragments: the

more complex grains are fragments of lava, cinders, glass, etc., all thoroughly bound

into a complex aggregate.

ing the acid and basic ends of the classification scheme, but here also the

rocks of the andesite-diorite family are by all means the most numerous

and most widely distributed. The greater number of occurrences are

represented by members of this family belonging to intrusives that would

be classed normally as andesite porphyries; porphyrites of various kinds

and diorite porphyries. The minor variations represented by these rocks

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 31

would probably include all of the habits known to this family. In addi-

tion to the porphyries of this family composition, there are less common

occurrences of felsite, quartz porphyry, granite porphyry and_ basalt

porphyry. In surface flows there is, besides the andesites, an occasional

amygdaloidal basalt, but so far as observed there was no rhyolite or other

very acid surface type. Among the massive larger intrusives, the com-

monest and most abundant type is a granite porphyry or granite varying

in some parts to the composition of syenite. A massive rock of the na-

Fic. 8.—Photomicrograph of a typical weathered shale from Fajardo Playa, magnification

28 diameters

The dark field is made up of an extremely fine aggregate of earthy materials. The

white circular and irregular spots are entirely empty and constitute the porosity of the

rock. The circular forms of these yoids suggest that they represent former calcareous

content in the form of foraminifera, now completely removed by weathering.

ture of a diorite is also represented, as is a very coarse rock of the nature

of a giant granite. How these are related, how many intermediate

varieties there may be and whether this variation represents magmatic

differentiation within a single mass or instead different units of intru-

sion, has not yet been determined in enough detail to make a positive

statement. But in at least two cases where these large masses were seen,

ANNALS NEW YORK ACADEMY OF SCIENCES

the hasty examination given to them leads to the belief that differentia-

tion effects can be traced.

The closest relatives of the igneous rocks are the sediments, and be-

cause of the fact that the material constituting these sediments has been

furnished by the volcanic fragmenial supply in large part, their character

and makeup is in many cases not strikingly different from the ashes and

finer tuffs. They do, however, represent an additional assorting, an ad-

ditional weathering and an additional opportunity for intermixture of

Fic. 9.—Photomicrograph of a foraminiferal shale from the Bayamon-Comerio road,

magnification 28 diamciers

The dark areas are chiefly earthy aggregates of very fine texture: the whitish areas

are calcareous spots which in many cases still preserve the forms of foraminifera. It is

the removal of such materials from the shales that is believed to account for their

porosity as seen in weathered outcrops.

materials from different sources and of organic material developing at the

same time. These conditions give a great range of composition and

mineral makeup to the shales and sandstones and they merge by imper-

ceptible gradations from simple tufaceous or arkosic sediments to eal-

careous rocks or even to fairly pure limestones. The common source of

the caleareous element in these rocks is from an intermixture of fora-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 33

miniferal matters representing organic growths accompanying the accu-

mulation of the deposit. |

The limestone members on the one side representing almost pure or-

ganic accumulations, and the conglomerates on the other representing the

simpler destructive volcanic fragmental matters, give-the range between

which an exceedingly great variety of sediments are represented.

The shales of the younger series, represented by the Lares and the

Juana Diaz shales, are more strictly detrital and of true erosional and

Fic. 10.—Photomicrograph of the San Juan formation indurated dune sand, magnification

28 diameters

The clearest grains are simple mineral fragments; the dark ones and the grayish ones

with internal structure are fragments of calcareous organic growths. The grayish matrix

is a secondary binding material of calcitic composition, in this case practically filling

the interstitial spaces.

destructive origin, and in places they contain lignitic material which

suggests different physical conditions. This shows, however, in their

upper layers an increasing amount of organic content also, and finally

are succeeded by limestones of wholly organic makeup. The failure of

volcanic activity during and subsequent to that time gave no opportunity

for the amount of intermixture that is seen in the shales of the older

series, so that as a result the younger series of rocks is petrographically

34 ANNALS NEW YORK ACADEMY OF SCIENCES

more simple and less modified. The organic content is of greater variety,

however, and because of the reef-building tendency the primary structure

is more varied than is seen in the limestones and shales of the pre-Ter-

tiary representatives.

The most striking petrographic type is, probably, the solidified dune

sand making up the San Juan formation. Its uniformity of grain,

strong cross-bedded character, porous habit, together with its rather sur-

prising stability, make it an object of some considerable interest. Sev- —

eral of these classes of rocks, therefore, represent petrographic series of

unusual range and variety, and because of their perfection of develop-

ment would seem to. warrant detailed study.

There are no foliated metamorphic rocks so far as yet seen in Porto

Rico. One specimen of such rock, a mica schist, was shown to the writer

as having come from the Portuguese river not far from Ponce, but a hur-

ried reconnoissance in the vicinity failed to uncover anything even re-

sembling it. The specimen probably does not belong to the rocks of

Porto Rico. No evidence whatever has been seen of conditions that

would be expected where such rocks occur. |

The most profoundly modified rock observed is a massive serpentine.

Such material was seen at two places by this party, one near Yauco and

another on the road to Comerio, and the same type is reported by Pro-

fessor Crampton on a much larger scale near Maricao. But they are al!

simple petrographic cases after all, being ordinary intrusive units of

heavier ferro-magnesian content than the average which have been heay-

ily altered, especially by hydration processes, to the present condition.

Genetically and historically, the serpentines are not materially different

from the other intrusive bodies.

Depth of Decay

Alteration has affected the rocks at most points to considerable depth,

but in spite of this there are plenty of outcrops, and along the roads there

are many cuts exposing fresh rock. The stream beds are strewn with

fresh pebbles and boulders. Although decay obscures the character of

the rock in most of the outcrops, the structure is usually fairly well pre-

served, and in most cases enough can be seen to enable one to determine

the formational habit.

The most striking thing about many of these badly decayed outcrops

and cuts is the remarkable way they stand against destruction or removal

by ordinary weathering and erosion agents. At many points, road cuts

are made, with side walls absolutely vertical, through wholly decayed

rock material, that stand for years without crumbling down. Embank-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 35

ments made of earth along roads and trails behave in the same way and

one is continually surprised at the steepness of such slopes and their ap-

parent stability. Slopes of 40° are not rare on hillsides that are culti-

vated, and one occasionally crosses divides that are mere knife edges with

slopes of this kind on both sides. Such stability of the soil is a great

factor in preserving the agricultural productivity of Porto Rico and in

the distribution of its agricultural industries over so much of the in-

terior area.

There seem to be three factors of large consequence in this stability of

the soil mantle. One is the clinging character of some of the vegetation

which tends to bind the soil together; another is the small range of tem-

perature variation which reduces disintegration or disruption tendencies

to a minimum; and still another is the low content of inert or refractory

materials, such as quartz, in the rocks whose destruction has furnished

the soils; all of which factors favor the making of especially tenacious

soil. Most of the soils are for the same reason exceedingly difficult to

cultivate. As a direct consequence of this soil behavior and climatic

control, there is comparatively little dust formed in Porto Rico. This is

especially noticeable on the roads, where one is almost never in the least

troubled by it.

In two districts, one on the north fringe of El Yunque along the

Sabana River and the other on the north side of the divide near Adjuntas,

deposits were seen which suggest glacial action. Very large boulders are

stranded in positions where it is difficult to account for them by ordinary

erosion means, but too little study was given and too little evidence is at

hand to warrant a more definite statement.

STRUCTURAL FEATURES

Most of the structural features represented in the geology are men-

tioned in connection with descriptions in other sections of this report.

This is especially true of such structures as may be regarded as essen-

tially primary. including the interbedded and intrusive forms of various

kinds. There is no doubt but that the most prominent structural com-

bination in Porto Rican geology is represented by the succession of inter-

bedded sedimentary beds and tuffs, cut by or interleaved with intrusive:

in the form of sills, dikes or irregular stringers. The combination occurs

in great variety as to succession, relative amounts or proportions, quality

of material, present condition and relations to other members, but in

spite of these variables the structural feature is essentially the same and

has the same meaning everywhere.

36 ANNALS NEW YORK ACADEMY OF SCIENCES

Igneous Structures

The most striking thing in connection with the structure is the re-

markable uniformity of the sills and their close resemblance on that ac-

count, after weathering, to the fragmental beds with which they are

associated. ‘The small amount of metamorphosing influence that they

seem to have had, also adds to the difficulty. In some cases, however, a

transgressive intrusion has disturbed the adjacent beds a great deal in a

mechanical way.

In the average case, it is judged that the intruded magma has neither

penetrated the materials of the adjacent beds to a noticeable amount, nor

Fic. 11.—Shales and ash beds cut by a large irregular dike and sill

The dike is shown at the location of the standing figure and the sill extends upward

to the left between the plainly bedded layers. Both the dike and the sill are crowded

with fragmental material to an extent that makes the intrusive have more the appear-

ance of a volcanic fragmental than a true intrusive.

has it absorbed or incorporated a great deal of such material. But in a

few cases where structural relations were indisputable it was equally

clear that the intrusives, both transgressive and concordant, were liter-

ally choked with foreign fragmental matter, making them resemble the

real tuffs so closely that it is doubtful whether the difference would have

been detected except for the clearness of the intrusive relation. Such

occurrences suggest that there may well be many other apparently frag-

mental interbeds that are in reality fragment-clogged intrusives. On ac-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 374%

count of the great abundance of the fragmental matter, it does not seem

possible that these intrusive masses could penetrate in that condition to

some of the positions where we now find them. It is more likely that a

rather fluid magma has penetrated some unusually porous fragmental

bed forming a matrix for it, perhaps also spreading it somewhat, and

then in breaking across to another bed, in some cases it was still mobile

enough to drag the mixture along into the larger transgressive structures.

This idea is somewhat supported by the finding of a conglomerate bed,

near K-86 on the military road west of Aibonito, impregnated with an

igneous matrix in essentially this same manner. In addition to these

forms, there are numerous larger intrusive masses, the largest of which

deserve a special name. I see no objection to calling them bosses. The

two largest occur, one between Caguas and the Caribbean sea toward the

south and southeast, and the other between Jayuya and Utuado on the

north side of the divide.

Volcamc Vent Complexes

A special igneous structure that has not been referred to except inci-

dentally is that composite of disturbed structures which represents the

location of old volcanic vents. They are essentially a complex of irreg-

ular intrusive units cutting and including masses and aggregates of

various fragmental and sedimentary types in a mixed relation. In the

clearest cases, such a complex suddenly takes the place of a formation of

apparent promise of continuity such as a series of sedimentary beds, and

after an interval these beds are again found continuing as before. For

example, the Coamo limestone formation is abruptly cut off and its place

is taken for a mile or more by one of these igneous complexes, the lime-

stone continuing on the other side again. The Jacaguas reservoir, just

above Juana Diaz, les in one of these old voleanic-vent complexes, occu-

pying, however, only a portion of the area. Another such case is repre-

sented by the complex cutting the great conglomerate beds on the

military road at about K-87—88 west of Aibonito. Another is believed

to be represented by the very striking basin-like area crossed by the road

between San German and Hormigueros. This one is now represented

by a very smooth plain five or six miles across surrounded on all sides by

more hilly country. The same conditions are undoubtedly indicated by

the extremely complex structures seen on the Descalabrado river two

miles below, south of the military road. Some of these mark the sites of

ancient craters, clogged or choked with fragmental and intruding ma-

terials.

38 ANNALS NEW YORK ACADEMY OF SCIENCES

Folding

Most of the rock formations representing original bedded types have

been more or less tilted or otherwise do not now have their original atti-

tude. Those belonging to the vounger series, the Tertiary limestones

and shales on the north side of the island, are comparatively little dis-

turbed, and in some cases do not have a very different dip in spite of the

fact that they have changed very much in position with respect to sea

level. On the south side of the island, however, even these later beds are

in many places tilted at a higher angle than they had in the beginning

Fig. 12.—Overturned fold and crush zone in finely bedded shales on the Jayuya road near

the summit of the range

and occasionally show high angles and even gentle folds. This condition

may be seen on the Jacaguas River, near Juana Diaz, better than at most

places, but similar conditions are indicated by the relations at certain

points farther west. This condition on the south side of the island indi-

cates more extensive and more violent dynamic disturbance on this side,

which is further supported by the presence of faults cutting and affecting

the Tertiary series on its present inner margin.

The older series, the pre-Tertiary, is still more profoundly affected

and, in almost all districts, shale and ash beds may be found standing at

high angles, in many cases practically vertical, and in occasional instances -

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 39

crumpled and overturned in a most complicated manner. Minor fold

structures of this kind are especially prominent in the higher ranges, for

example, along the Jayuya road along the divide opposite station K-24

of the Ponce-Arecibo road. At such a place may be found as compli-

cated structural features of this kind as is usually present in any folded

mountain region. The high angle at which such beds stand at many

other points leads to the belief that similar complexities characterize a

great many of the districts occupying the higher mountain portion of the

island as well as some of the marginal areas, but the great amount of

erosion and the very limited exposures at most points tend to obscure

some of these details. The complex way in which the igneous intrusive

masses cut all of these formations also tends to obscure and modify and

further complicate the simpler fold structures so that it is not always

possible to properly credit the disturbed attitude. On account also of the

‘fact that the total quantity of injected or intruded materials, including

dikes, sills and bosses, is exceedingly large and must have caused exten-

sive disturbances by reason of the displacement produced by the occupa-

tion by these intrusive masses, it is hkely that much of the observed ab-

normal attitude of the bedded rocks may be due to this cause rather than

to regional folding of a simpler sort. It is fair to say, however, that a

sufficient amount of data is not yet available to draw general conclusions

as to the meaning of the fold structures in Porto Rico. The striking

thing is that all of the older formations are disturbed and that their

position and attitude, even along the margins of the island, indicate that

the region affected by these movements was more extensive than the

present land area.

Faulting

There are many evidences of faulting on a small scale, in some of which

the displacements can be measured. But in most cases the direct evi-

dence lies chiefly in the existence of crush zones, slickensided walls and

abrupt changes of rock type; there is no opportunity, on account of the

general rock complexity, to secure quantitative data. Judging from the

difficulty in tracing certain formations between districts where they have

been identified, it is probable that there are occasional faults of large dis-

placement. Numerous crush zones were seen on the Comerio road espe-

cially, but in this case also no system was discernible from the few meas-

urements available.

The most prominent fault, in its effect upon present features, is the

one now marking the inner margin of the younger series of chalky lime-

stones and shales constituting the coastal belt along the south side of the

40 ANNALS NEW YORK ACADEMY OF SCIENCES

island from Juana Diaz past Ponce at a short distance to the north, eross-

ing the Ponce-Arecibo road at K-4.8, and thence westware, crossing the

Ponce-Penuelas road at K-10. This is the only large fault actually ob-

served that is necessarily of recent age, although a few others are inferred.

abruptly cut off by it. The older rocks of the pre-Tertiary are lifted with

respect to the younger series forming the present coastal margin wherever

this fault has been seen. It has been traced by us from Juana Diaz to

the vicinity of Penelas, a distance of about 12 miles. What becomes of

Fic. 13.—Crumpled shales as seen along the Jayuya road near the summit of the range

it at either end is not yet determined, but it is believed to extend much

farther in both directions.

The physiographic habit of the island as a whole tends to support the

view that the fundamental structural form is that of a large fault block,

with the principal fault displacement and uplift along or near the south-

erly margin, tilting the whole mass gently northward. If this disturb-

ance took place. as seems to be indicated by the fault described, in very

late Tertiary time, accompanying the emergence from the sea, it would

account for the abnormalities of Tertiary rock distribution as well as the

unsymmetrical position of the main drainage divide. Im any case, how-

ever, the fault block structure is a very late development and is superim-

posed on the other more complex and older structures of the mass.

BERKLEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 44

The island is comparatively abruptly terminated at both the east and

west ends. ‘The younger limestone margin, which is fairly continuous

along the north coast and extends along about half of the south coast, is

wholly wanting at the east end and is also absent at the west end except

at the corners. At Fajardo, at the east end and at Rincon, at the west

end, for example, the older complex bedded rocks continue to the shore

line. In the uplifting of the present island mass, it would therefore ap-

pear that breaks occurred at both ends. The included mass is therefore

probably bounded roughly on three sides by faults, the east, west and

south, and is as a block tilted gently to the north.

That there is, besides, considerable differential movement accompany-

ing the uplift and disturbance, is indicated by the warping of the erosion

plain lying beneath the younger series, the Eocene peneplain, which

stands essentially at sea level in the vicinity of Loiza and more than a

thousand feet above it at Lares. This difference is accompanied by a

much wider belt of these later limestones also in the region about Lares

than elsewhere. Such warping need not of course be confined to the last

movement; it may have accompanied the depression in eatly Tertiary

time, permitting, as is indicated by deposits, very marked differences in

the development and encroachment of the organic accumulations.

Large Structural Groups

Where rock formations or field units are so numerous and so closely

related and so complex in primary structure if taken in detail, it is ad-

visable to combine them into fewer more generalized groups. A first step

of a very general sort, but im all respects sound, has been taken in recog-

nizing and using the terms “Younger Series” and “Older Series” in this

paper. An additional step has been suggested in recognizing certain

smaller associations under the terms San Juan Formation, Arecibo For-

mation, Coamo Limestone Formation, Juana Diaz shales and marls,

Fajardo shales, Sierra de Cayey tuffs, Ponce chalky limestone, etc., but

these are for the most part local designations, some of which may well be

expected to become unnecessary after complete correlation is established.

A good structural basis for sound subdivision of this sort is not yet

worked out.

Unconformities

There is only one marked unconformity in the structure of the island.

This is between the younger and the older series. It measures the break

in the sedimentary succession represented by the erosion interval during

which this mountain mass, now represented by Porto Rico, was reduced

42 ANNALS NEW YORK ACADEMY OF SCIENCES

to a comparatively monotonous surface for the most part at least near to

the sea level. The time interval need not have been of very great geo-

logic value, but it represents the time between the last violent outbursts

of volcanic eruptive activity. occurring near the close of the Cretaceous,

and the beginning of simple sedimentation and limestone reef develop-

ment and other organic accumulations in the early Tertiary. This un-

conformity is very pronounced along the northern margin of the island

wherever the two principal series of formations are well developed. This

is not easily seen on the south side, but the relative complexities of atti-

tude of the two series, together with their position, emphasizes the same

fundamental relation. This break in sedimentation is not anywhere

marked by a development of a basal conglomerate. In some places the

new series is Inaugurated by the development of shales, part of which are

hgnitic, indicating land conditions, but in other places such beds are

entirely wanting and the upturned eroded members of the older series

are followed abruptly by limestones of the reef type. The first type of

succession is illustrated in the vicinity of San Sebastian and Lares and

the latter type of abrupt limestone succession by the conditions seen on

the Arecibo River. It is entirely lhkely that the time value of this break

is not everywhere the same. Probably the districts in which shale beds

are developed saw the beginning of sedimentary deposition at an earlier

period than those in which the shales are entirely lacking. It will be pos-

sible to work out these historical and structural differences with further

study of the content and distribution.

A very extensive development of conglomerates in the region immedi-

ately west of Aibonito and smaller occurrences at several other points,

especially on the Comerio road south of Bayamon and also near the

military road in the vicinity of La Muda, have a suggestion of the possi-

bility of a rather important structural break, but there is thus far no

conclusive evidence of the presence of any real unconformity.

Veins

Quartz veins are not prominently developed. There are occasional

quartz stringers and in a few places they are abundant enough to make

a sort of net work, but in no instance was a large persistent fissure vein

seen. In some cases the stringers of quartz carry values in gold, and in

all probability they are the source of the placer gold known to exist in

certain districts. So far as observations have been made, there is no par-

ticular system represented in the vein occurrences. Questions connected

with this subject together with mineralization and possible value as

mineral resources should be made a special study.

BIRKEY, GHOLOGICAL RECONNOISSANCE OF PORTO RICO 48

Minor Structures

Although there is extensive development of sedimentary formations

which have been subjected to much disturbance, there is comparatively

little structure of a minor sort that seems to deserve such discussion in

this description. ‘Two, however, that seem to have special significance

connected with the origin of the particular beds which have been found

are (a) a peculiar crumpled, enterolithic structure seen in one of the ash

beds near Guayama, and (0b) the wind-assorted cross-bedding structure

of the old dune sands of the San Juan formation at Arecibo.

Enterolithic Structure-——The enterolithic structure noted in the ash

beds, on account of the thinness of the bed,—about eighteen inches,—and

the simplicity of the associated structure,—simple tilted beds,—leads one

to believe that the structure is essentially primary rather than of subse-

quent dynamic origin. Its appearance is perfectly consistent with the

explanation that it is preserved from the time of deposition and its be-

havior at that time as a small mud flow. It is a structure such as might

be formed by slumping movement of a soft layer. It should be expected

that there would be frequent behavior of this kind in the accumulation

of such extensive beds of ashy materials, which must in some cases have

been deposited under conditions that would make slumping movements

possible, but it is not to be expected that material of this kind would in

most cases be capable of preserving any of these primary movements. In

the case noted, the quality of the interior makeup of the bed seems to

have been more favorable to such preservation. It is the only case where

such an observation was made.

Double Cross-bedding—The cross-bedding structure belonging to the

San Juan formation is a prominent feature wherever these ancient dune

sands are preserved. A great prevalence of steeply inclined minor struc-

tures is crossed by fewer nearly horizontal ones. Measurements made on

sea-cliffs a short distance west of Arecibo, where this rock is very promi-

nently developed, gave dips of 30 to 33 degrees repeatedly. A series of

these is abruptly terminated by a more nearly horizontal bedding for a

comparatively short distance and the whole structure is repeated. The

layers with this kind of structure are prevailingly one-half to two feet

thick and no ripple marks were seen on any of the beds examined. The

eross-bedding structure in this case dips always to the west or southwest,

and the average strike of the principal beds is about north 30 degrees west.

This is consistent with a wind direction not very different from the pres-

ent prevailing winds. Very strong structural development of this kind is

also to be seen in the city of San Juan at the promontory on which the

AA ANNALS NEW YORK ACADEMY OF SCIENCES

Morro is built, but measurements of orientation were not taken there.

An occurrence of this rock immediately to the east of Arecibo, a short

distance south of the lighthouse, showed structures of this kind on a

much larger scale than was seen elsewhere, and with an especially imter-

esting combination structure. The principal or stronger divisions are

widely separated and le nearly horizontal. A comparatively small bed

lying in this position was almost unconsolidated, but those strongly cross-

bedded immediately above as well as those below were compact enough to

stand in a vertical cliff 30 to 40 feet high. The chief interest attaches

Fic. 14.—Cliff of the San Juan formation south of the lighthouse at Arecibo

The prominent cross-bedding, extending throughout the upper thirty feet of the cliff,

is well shown, together with a less strongly marked horizontal structure crossing the

same beds. The prominent break near the base is made by a layer of sand which is very

poorly consolidated.

to the strongly cross-bedded portion forming the upper twenty feet or

more of the exposed cliff. The cross-bedding structure itself extends

without interruption through a much greater vertical range than in any

other outcrop examined, but its attitude and dip were not markedly dif-

ferent; the feature that was strikingly different from the structure seen

anywhere else was introduced by the presence of less pronounced but still

very plainly marked horizontal structures, making an interpenetrating

mesh-like arrangement in the face of the cliff. This can be seen strongly

enough to show in a photograph even at a distance of 100 feet. It is evi-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 45

dent that some peculiar condition has been in control in the development

of this compound structure. There is no reason to assume any difference

of source or origin for the steeply inclined structures, or cross-bedding

structure, from that assumed for all of the other occurrences of the San

Juan formation. It is apparently a perfectly normal product of the as-

sorting action and deposition of material under the work of the wind.

But under normal conditions it would not happen that a second structure

in a horizontal position should be repeatedly developed crossing the well

marked cross-beds so that the whole complex combination should be de-

veloped on such a scale as is seen in this case. From the nature of the de-

posit and the conditions in which it may well be assumed to have been

formed—that is, at or near sea level in the vicinity of or bordering upon

standing bodies of water—one would be inclined to favor the explanation

that the accumulating cross-bedded sands fell into or rolled into a body

of water which had a tendency to attack the newly deposited material and

to bind the grains together. The difficulty with such an explanation is

in the fact that the horizontal structure is repeated at small intervals

practically throughout the deposit, and apparently without disturbing

the primary depositional structure at all. It would seem quite unlikely

- that loose matters of this kind, falling into or rolling into a body of water

of an open surface sort, should maintain or preserve the primary struc-

ture so well. Perhaps it is more logical, in view of all the features, to

connect the development of this structure which seems, from its slight

influence or modification of the cross-bedding, to be wholly secondary

with the subsidence of the coast which is shown to have been one of the

late events in the geological history. As subsidence progressed, perhaps

somewhat irregularly, it would happen that the ground water level would

rise correspondingly high in beds that were passing below sea level. At

the surface of this ground water level the tendency would be to accomplish

a binding of the granular materials together. Both above and below

the ground water level there would probably not be so strong a tendency

to develop this binding. With the next step im the progress of subsidence,

another streak or indurated zone would be established and these have

been repeated throughout the whole formation during the whole period

of subsidence. An action of this kind would not tend to disturb the pri-

mary structure at all; it would on the contrary tend to preserve it or

make it less destructible because of the improved induration. But it

could, in addition, develop a succession of secondary structures through-

out the whole mass which, if the binding is fairly substantial, might rival

the primary structure in prominence when exposed to subsequent de-

structive attack. It is possible that such a succession of horizontal struc-

46 ANNALS NEW YORK ACADEMY OF SCIENCES

tures could be developed even under a perfectly continuous but very slow

subsidence movement by reason of the natural seasonal ground-water

fluctuation. From this point of view, the range between two succeeding

Fic. 15.—Detail of the double structure in the San Juan formation at Arecibo

This photograph was taken at the same point as the one shown in Figure 14 to bring

out the horizontal structure crossing the inclined layers. There is no doubt whatever

that the dark layer of less consolidated sand in the lower third of the photograph is a

primary bedding structure, but the horizontal marks crossing the inclined layers in the

upper part of the view are believed to be of secondary origin.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 4%

horizontal markings would measure the fluctuation range of the ground

water, the harder zones representing in that case the more persistent

upper level for each succeeding depression position.

The cross-bedding structure shows as plainly as it does on exposed sur-

faces also because of the fact that certain streaks are more perfectly in-

durated than the intervening ones, and the objection might well be raised

that a history of the kind suggested above would not be expected to de-

velop such a difference of induration in layers inclined at such high

angles to the horizontal. As a matter of fact, however, these sands are

not simple in their makeup. They are in large part fragments of organic

material and complete shells of small organisms of a calcareous nature

and the primary cross-bedding structure represents an assorting action

on this mixture of silicate and carbonate mineral material. It so hap-

pens, therefore, that the successive structural units are not necessarily of

the same mineral proportions, and in the process of induration, or of

binding the grains together, certain streaks yield more readily to this

influence and develop greater solidity and resistance to destruction. It

happens, furthermore, from a variety of rather unusual primary condi-

tions and secondary influences that both a primary and secondary struc-

ture of unusual prominence and peculiar association are developed in the

San Juan formation.

SPECIAL RELIEF FEATURES

Playas

The flat areas along the coastal margin which are known as playas are

all developed at the mouths of rivers and are essentially alluvial deposits

of floodplain and delta type. In most cases they seem to occupy areas

that must formerly have been embayments in the coast. This develop-

ment is most striking, for example, at the mouth of the Arecibo and of

the La Plata and Loiza rivers. In some cases, however, there is no

marked embayment and the deposit is more strictly marginal, such, for

example, as the Fajardo Playa at the east end of the island and others

on the south coast.

Promontories

In addition to the embayments and playas, there are, occasionally, in

the intervening spaces, promontories where the rock formations extend

into the sea and terminate in cliff forms. These are neither numerous

nor are they confined to any particular portion of the island or to any

rock formation. They are represented by the most recent of all of the

48 ANNALS NEW YORK ACADEMY OF SCIENCES

formations, essentially a silicified dune deposit such as that at San Juan,

also by Tertiary limestone reefs, such-as that at Quebradillas and at

Guanica, or by the still older igneous and clastic series, such as that at

Anasco, or by massive intrusives of a strictly igneous habit, such as that

at Maunabo. It would appear from this that the former outline of the

island must have been more irregular than it is at present and that the

distribution of marginal formations is also not as regular as has been

represented in earlier reports.

Fic. 16.—Playa plain and marginal terrace

View of the Playa plain (foreground), the marginal terrace (middle field) and the

mountainous divide formed by the Sierra de Cayey as seen from the ‘Central Machete’

near Guayama. This terrace bevels across the upturned edges of shales, ashes and intru-

Sives of the older series and is probably due to marine cutting.

Terraces

At many places on both sides of the island there are comparatively

smooth tracts having the appearance of bordering shelves which represent

true terraces. Their location along the sea margin and the compara-

tively insignificant development of similar benches along the streams

lead one to believe that they have an origin connected with the wave

action and attack of the sea.. This interpretation is supported by the

presence of these terraces along the coastal margin where stream action

would not seem to have been able to reach. In any case, the presence of

such terraces, which stand from 100 to 200 feet above the present sea

level, indicate a former more submerged condition, so that the sea or

streams, or both combined, were able to attempt base-leveling at that ele-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 49

vation. The presence of great quantities of roughly assorted gravels

clogging some of the valleys of the southerly side of the island tends to

support the same general conclusion. The bearing of these features on

the geological history of Porto Rico will be taken up at another point.

Cuestas

Both on the north side, for nearly the whole length of the island, and

on the south side, over the westerly half, there is a bordering belt of

limestone and associated beds that have been developed on an eroded sur-

Fig. 17.—Structure beneath the marginal terraces

Strongly bedded ash together with associated shales cut by small dikes forming a part

of the terrace near Guayama. These rocks belong to the older series and dip into or

toward the mountains rather than toward the sea.

face which beveled across the more complex structures of the older series

of formations that formerly made up the mass of the island. These

limestone beds are several hundred feet in thickness and dip gently

toward the sea. On the inner margin of their present extent toward the

interior, especially along the north side of the island, they are abruptly

terminated in a very irregular line of modified cliff forms facing toward

the prevailingly smoother and lower ground for some distance toward

the interior. For the most part, this limestone margin is exceedingly

rugged and broken. The width of the belt with this rugged character

50 ANNALS NEW YORK ACADEMY OF SCIENCES

varies very much in different parts of the field. Its most striking devel-

opment is in the district extending from Tao Alto to Aguadilla. In the

district extending eastward from San Juan and also in some of the areas

on the south side, this margin is very much broken and so obscure in

some parts as to escape detection. In its best development. however, it

is a typical cuesta, formed in the usual manner by the erosion of a for-

mation representing a recently uplifted coastal series: ‘The series of

formations involved formerly extended inland very much farther than

they do now. Only the outer margin remains from the erosional de-

struction of a series of beds and reefs that in former times covered a

Fig. 18.—IJnner lowland near Bayamon

View looking north from the Bayamon-Comerio road toward San Juan, showing the

monotonous features of the lowland belt in the foreground and the comparatively promi-

nent hill remnants of the Tertiary formation cuesta in the background.

large portion of the island. -The road running from Aguadilla to Moca,

San Sebastian and Lares extends for practically the whole distance, after

leaving the coast, along the inner lowland at the foot of this cuesta or

along the cliff forming the inface. The same features characterize the

surface topography as far east as Corozal. This feature is much less

pronounced on the south side of the island.

Peneplain

Beneath the limestones constituting the cuesta and representing the

Tertiary series there are, in numerous places, traces of a former plain

that represented the results of erosion on rocks that had a complex struc-

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 51

ture. Occasional profiles of more distant ridges also show a sky line that

suggests the former existence of such a plain, and in favorable localities

it can be traced directly to the foot of the limestone cuesta. Occasional

traces are also seen on more mountainous tracts, especially at the west

end of the island, near Rincon and in the vicinity of Mayaguez. At the

latter place, these remnants of the old plain are called mesas. It is the

judgment of the writer that these all belong to a single base-leveling sur-

face or marine-cut platform formed in the period just preceding the

development of the Tertiary limestone series. Judging from beds devel-

oped immediately upon this surface, it must have been completed in early

Fig. 19.—Haystack (pepino) hills

A characteristic view, showing the small soil-covered flats and associated haystack

hills found in the region of the Arecibo formation. Photograph taken on the road be-

tween Arecibo and Barceloneta.

Kocene time, and perhaps was even largely developed in pre-Tertiary

time. It may be referred to as the early Tertiary base level or peneplain.

There are many minor features giving variety to the surface relief

which depend for their particular relations and character on underground

structures which are as yet imperfectly understood.

Haystack Hills

The most striking topographic feature of the whole island is the re-

markable development of small isolated or grouped rugged hills usually

rising abruptly above adjacent smooth flat soil-covered areas at various

levels throughout a broad marginal belt along the north coast, west of

59 ANNALS NEW YORK ACADEMY OF SCIENCES

San Juan. They constitute a feature so unusual that even the un-

trained casual visitor is impressed with them.

This feature has been referred to before in connection with the de-

scription of the “Younger series” of rocks, especially the Arecibo reef

limestone formation. In spite of the unusual appearance presented by

this distribution of “haystack” hills and intervening flats, their origin is

judged to be comparatively simple. The active agents and processes

have been the same as those at work on all other parts of the island, but

the results differ because of the fundamental difference of material and

Fig. 20.—Care structure in the haystack (pepino) hills

Near view of the limestone hills forming the margins of the small cultivated flats in

the typical haystack hills district. This view shows the cavernous nature of the lime-

stone forming these hills, a structure that is regarded as the most significant feature

and probably the largest factor in the development of these peculiar relief forms.

structure. Nowhere are these features developed except where the later

reef limestones are the underlying bed-rock formation.

The essential steps in the development of these forms are the follow-

ing:

The reef limestones are not uniform in composition or structure.

They have more or less intermixture of earthy matters which are distrib-

uted irregularly, but chiefly at certain horizons, as more earthy or shaly

beds of no very great lateral or vertical extent. Such conditions are re-

BERKEY, GHOLOGICAL RECONNOISSANCE OF PORTO RICO 58

peated at occasional intervals in successive horizons. As such a series is

lifted above sea level and subjected to ordinary erosion and weathering,

the tendency is, (a) for the purer and more massive reef limestones to be

attacked by the solvent action of percolating water with a development of

underground channels, porous rock condition and actual caverns, (b) for

the more earthy layers to resist and limit such action at the levels where

this matter is present in sufficient abundance, with a development of

residuary material. As this action progresses toward maturity, many of

the larger caves collapse and sink holes are thus formed. With still fur-

ther development, the sink holes merge into each other in local areas

where solution has been most active, the earthy debris forms a soil in the

bottom corresponding in level with the first important earthy layer, and

adjacent remnants of the limestone reef stand out as sharp rugged hills

separated by irregular notches that represent other smaller collapsed

eaves. The result of such action and conditions, finally, is the numerous

“haystack” hills standing on flat soil-covered areas or surrounding such

areas as if they were just set down as bunches on this surface. This re-

lation is repeated at different levels throughout the belt from San Juan

to Aguadilla, but the most striking developments are local, apparently

where the structural relations are just right, and may be seen best be-

tween Tao Alto and Arecibo, especially in the vicinity of Manati and

Vega Alta.

It was at first thought that former subsidence levels might have some-

thing to do with establishing the level tracts, but the observation that

these tracts stand at. very different levels in immediately adjacent dis-

tricts together with recognition of the structural difference, lead us to

give credit to the primary structural character of the formation itself as

the controlling factor in the present distribution. According to this ex-

planation, these hills are mere remnants left from solution attack on a

reef limestone, the depressions between them representing collapsed cay-

erns, the walls of which may still be seen on the sides of some of the more

rugged hills, and the surrounding or intervening tracts are soil-covered

and level, chiefly because of the accumulation of earthy material, left be-

hind after removal of the overlying reef, now halted in its reduction at

the first important less soluble beds.

MINERAL RESOURCES ©

An examination of specimens of minerals and ores in the hands of

local prospectors and residents interested in developing mineral resources,

®A good list or tabulation of the mineral occurrences of Porto Rico may be found in

the article by H. C. B. Nitze listed at the close of this paper.

54. ANNALS NEW YORK ACADEMY OF SCIENCES

together with observations made personally, shows that there is consider-

able range of minerals and ores. It appears also that considerable atten-

tion has been given in a few cases to local development. ‘There is large

variety shown in a collection of this material and in some cases the speci-

mens exhibited look very promising indeed. But there is almost no

reliable information touching the quantity or the exact relations or esti-

mates of possible profitable development. It can be said, without danger

of contradiction, that none of the developments so far undertaken looking

toward the systematic mining have proven profitable.

Gold

Only one enterprise of this kind seems to furnish any production, and

this is the placer mining for gold. Gold washing has been practiced

from the early Spanish occupancy to the present time, and it is not at

all a rare thing to see several men digging in the stream gravels for the

“nay dirt” and panning out the gold. This is done in all cases on a very

small scale and with the aid of the simplest equipment, and the returns

appear to be very moderate. It is claimed that in former times a much

more elaborate system of working such deposits was in operation under

the Spanish regime, and, according to historical statements, they were

‘ considered profitable. More recently, there has been at least one attempt

near Corozal to develop this kind of ground by the use of modern ap-

pliances, but the plant has been allowed to go to entire ruin. The only

places where actual placer washing was seen in progress was three miles

south of Corozal and on the Sabana river near Luquillo. Near Corozal,

also, some work has been done in an attempt to discover the veins or lode

which may have furnished the placer gold. There are several pits,

trenches and shafts, in some of which quartzose stringers were seen which

appear to fulfill the requirements of a source of supply. Some free gold

was found in panning a little of the weathered material at one of these

spots. There is little doubt but that these veinlets or stringers, which

were numerous at one of the cuts, are in part the sources of the placer

gold of this locahty. But at no place examined was there to be seen any

“vein” of apparent consequence or any structure suggesting the course

or extent of the mineralization. Of course the rather mixed state repre-

sented by the residuary matter, seen almost everywhere at the surface,

does not lend itself readily to the tracing of veins, and it may therefore

happen that conditions would prove, after thorough exploration, to be

better than the first brief examination indicated. There are said to be

some old abandoned workings dating back to Spanish conquest times at

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 55

the same locality, but such evidences are very obscure and would seem at

best to have very little bearing on present prospects.

Copper

Some very good specimens of copper ores were seen in the possession of

Mr. Henry D. Sayre, of Corozal, who assured us that there were several

localities represented in the collection and that in no case had the real

value of the occurrence been determined. It was understood that exami-

nations have been made by engineers sent to Porto Rico in the interest of

American mining companies, and that some exploratory work has been

carried on by. Porto Rican companies or groups of individuals, but so far

as known there is at the present time no development work being con-

ducted, and the possible value of these deposits has not been thoroughly

proven.

Zinc, Lead and Silver

One prospect, on which several hundred feet of underground work has

been done, was visited at Barrio del Carme in the Sierra de Cayey, on

land owned by Pablo Vasques, several miles northwest of Guayama. The

country rock is chiefly andesitic tuffs cut by porphyritic intrusives. A

quartz vein carrying sulphides, pyrites, sphalerite, galenite and chalcopy-

rite has been followed and there is some ore on the dump. ‘The vein

varies from a mere streak to a width of two feet. The mineralization is

irregular and the values are said to be chiefly in lead and silver. The

second-class ore is essentially mineralized tuff. The exploratory work

has been done in large part on side slips and streaks quite outside of the

vein proper. In all of this side work, there was apparently no new min-

eral-bearing ground discovered. The first-grade ore is heavy and the dis-

tribution of values is not determined. There is no doubt of the existence

of a real vein or of the ore in this case, but there is need of more intelli-

_ gent exploratory development along the vein proper, together with a

study of the possibilities of separating the chief values by some sort of

milling operation, before a reliable conclusion could be reached as to pos-

sibility of working the deposit as a mine.

Iron

- One magnetic iron prospect was visited. This occurrence is on the

divide about ten kilometers west of Naguabo. It is reached by driving

eut on the road from Naguabo toward Torres to about this distance and

then taking saddle horses to the divide, a distance of about two kilometers

southward. On the expedition we were accompanied by Mr. Arturo

Gallardo, Jr., Alealde Municipal of Naguabo.

56 ANNALS NEW YORK ACADEMY OF SCIENCES

There are many surface bowlders of magnetic iron of fine quality in

this vicinity. A lhttle underground working is evident at one point but

this is now caved in. Surface observations, together with a few magnetic

observations, failed to show any very extensive deposit at that point, but

the quality appears to be good in iron content. The ore carries a little

copper and is associated with an igneous rock essentially andesitic in

composition. It could be traced with a fair degree of certainty about

fifty feet east and west just below the crest of the ridge. Considering the

associations at this place, it seems necessary to conclude that the ore is

igneous in origin and that it probably accompanies one of the porphyrite

intrusives. Other occurrences of similar ores were mentioned to us in

this same region, but none were visited.

Coal and O1l

There is no good ground for believing that valuable resources of these

products exist in Porto Rico. Some prospecting for them is carried on,

however, in a desultory way. The only basis for the hope of finding coal

is the occurrence of lignite and lignitic material with the shales lying at

the base of the younger series of rock formations, below the Arecibo lime-

stone member. JLignitic material was seen in these shales near Lares,

and similar or better material has been reported from near San Sebas-

tian. From what has been seen, there seems to be no promise of very

valuable deposits of this kind. The structure is simple and a very little

exploratory work done in a systematic manner would determine the prob-

able value of every occurrence known. There is no promise at all of such

content in the older series.

No oil indications were observed. The only formation to be considered

in investigating the prospect of oil is also the basal shales of the younger

series.

Limerock

A particularly porous, granular and uniform limerock is obtained from

the small island, Icacos, just off the northeast coast, and is used in sugar

refining at the Central, owned by the Bird brothers, at Fajardo. The rock

is organic, largely foraminiferal, and is probably structurally of the same

origin as the San Juan dune sand deposits,—comparatively recent. Such

materials are doubtless to be found in large amount, but not always so

pure and so uniform in quality and structure. Limestone suitable for

lime burning or for cement mixture is certainly not rare. Limestone of

a quality that would permit its use in structural work is also found at

some places, but apparently very little native stone is used.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO. 57

Guano

Bat guano is found in some of the caves in sufficient amount to be a

source of local fertilizer.

Road Metal

Several kinds of stone were seen used extensively in road improve-

ments. The particular variety used depends largely on the local supply,

but the most common are the Arecibo limestone and the massive syenite

and granite porphyry. There is no lack of these as well as other types

suitable for such use.

There are other mineral substances that will invite investigation, but

no-others came under the writer’s personal observation and no facts re-

garding them are in hand.

HOT SPRINGS

Thermal springs are known in the vicinity of Coamo. Judging from

their location and apparent relation to other physical features, they are

believed to lie along a fault weakness. The district is also one of com-

paratively late igneous activity, and this, coupled with the other factors,

leads to the suspicion that the springs are directly connected with the

dying igneous activity and may actually represent juvenile waters.

At Coamo Springs Hotel, one of these springs has been developed and

controlled for commercial purposes. The water is hot as it comes out of

the side of a small ravine and runs down over the slope, which is covered

to a moderate extent with deposit from these waters. The immediate bed

rock is not well exposed, but it is judged to be either a tuff or an igneous

complex and the field relations in the vicinity show that there is a thick

series of beds both above and below. No doubt critical field study could

determine the actual relations with considerable certainty.

An analysis of these waters, furnished by the proprietor of the springs

and made at the agricultural experiment station at Mayaguez, is as fol-

lows:

Fixed elements per litre of water

Biber sania seit... Beto l coe ae ae ek eek 0.01296

RE OTe HIN Mor s= semie ote ts. ct alt cb, creole Late 0.79902

SSELFPLODESSLT ESOC eae ae a a ee a en 0.52531

NCEA AOL ASST: oe oe es air ebm Scie stile even ewes se O.Q0082

UUM GE SOIT co ccbak fcc vey sce cdua Gace esbece’e 0.23054

0 LEE SUT 1 ERS AE ee a ee 0.08127

LSP IGT SUPES U1 ONS 00 hapa 2 ne Oe ee 0.03503

Carbonate of iron........ See hale d ea tciors ou Wd dete S Ab ace 0.01114

ee eas acre na) Sib. oie so Re hae view ee es 1.68559

58 ANNALS NEW YORK ACADEMY OF SCIENCES

Gases in solution per litre of water at 0° of temperature and 760 mm. of

pressure

Nitrogen: -. 223 «3 oes oie Oe a eee eee 13 ce. 740

OXY SOB. 2255 kn FRE Stee es oe he Ree ok ee 761

Sulphyd@ric Reise cee see ae ee ae ers 1 967

Potal Fo kas 2 eae ek tee 15 2468 -

HistTorRIcAL STATEMENT

A complete or even a reasonably full account of the geological history

of Porto Rico cannot be written at this stage of the investigations. Such

a statement is necessarily the end product or climax of the whole series

of studies that are proposed, but it may not be out of place to outline

some of the leading and most clearly marked steps as a rough sketch or a

preliminary attempt.

At the outset, it is well tc appreciate that the Island of Porto Rico is

geologically young. There are no traces, so far as known, of any of the

so-called ancient rocks. It is quite true, of course, that the older series

of formations is largely a volcanic complex whose exact age may never be -

accurately determined, but there is no occurrence of profoundly meta-

morphosed members or other evidences of great geologic age. Besides,

the series, complex as it is and difficult to group into suitable divisions as

it may be, undoubtedly forms a very closely related succession of minor

formational units whose uppermost members are determinable as to age

with reasonable accuracy. It would appear also from the nature of the

deposits and their structural relations that the accumulation must have

been, for the most part, a rapid process.

There is no good reason, so far as any of these facts are concerned, why

the whole of the “Older series” could not have been accumulated in a

single geologic period. ‘The fossil content of the upper members of this

series indicates that this period was the Cretaceous, as used in the broader

sense in geology. Whether or not the older members date back to an im-

mediately preceding time cannot yet be definitely stated, but whatever

there is, is clearly so closely associated with the Cretaceous beds that they

can all be treated as a single historical unit.

This earlier period is characterized by volcanic and other igneous activ-

ity on a very large scale. Beds were accumulated both above and below

sea level. There seem to have been oscillations of level accompanied by

recurrences of similar beds, and apparently much shifting of the supply

of materia] accompanied by great variation of character laterally. There

is good evidence that succeeding volcanic outbursts broke through these

beds at many places.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 59

An occasional more prominent change of conditions, more or less

clearly marked in the structural relations and character of material, may

possibly be used as a basis for epochal subdivision. It is quite clear, how-

ever, that there was no profound change of geologic control throughout

the whole of this earlier time,—-it was strictly a volcanic period. The

succession of disturbances by which it was affected is represented in part

by dynamic modifications of the nature of folding, crushing and faulting,

but this is probably an accompaniment of the more profound igneous

activities also, and need not be regarded as evidence of any strikingly

different causal process. If there were contributory causes of a regional

sort controlling the folding, they are essentially simply superimposed

upon or introduced into the larger, more profound and longer continued

igneous activities which prevailed both before and after that time.

This long period, characterized by great complexity of formational

development, including tuffs, agglomerates, conglomerates, shales, lime- ~

stones and immense numbers of intrusives of great variation in size, form

and composition, finally came to an end by the dying out of the volcanic

energy, and greater stability of the whole with respect to elevation and

subsidence was established. Erosion cut down the exposed formations,

the sea attacked the margins and in time most of the projecting moun-

tain mass was reduced near to base level, the sea encroached far onto the

former land areas and a new historical chapter was begun.

It is not possible to say, with the data in hand, that the entire island

was reduced to a peneplain, or perhaps a conoplain, but there is good evi-

dence, from the traces still left of former planation and from the dispo-

sition of the remnants of overlying formations still preserved, that the

greater part of the present area was worn down to base level and sub-

merged. ‘The process of base leveling was going on before the close of

igneous activity and it was continued long enough to bevel across rocks

of all sorts with marked success, but there is no necessity for regarding

it as a very long geological time.

As erosion proceeded, sediments were deposited unconformably around

the margins of the island of that time and perhaps also in some of the

marginal valleys above sea level. These constitute the earliest shale beds

of the “Younger series” and are believed to be of Eocene age. They are

at least early Tertiary. Where more simple marine conditions came into

control, as would happen when submergence or planation had masked or

destroyed the more elevated sources of supply, the deposits became almost

wholly reef limestones and shell limestones, with only minor amounts of

strictly detrital material irregularly distributed. This gave a succession

of somewhat irregular beds which are abundantly supplied with organic

60 ANNALS NEW YORK ACADEMY OF SCIENCES

remains and which bear evidence of the continued depression favorable

for the growth of these accumulations for a considerable part of Tertiary

time. ‘There is some suggestion in the relations shown in the eastern

portion of the island that this end was not wholly submerged and that

differential subsidence gave to this portion less prominent development

of the heavy, massive limestone beds.

In later Tertiary time there was marked reémergence from the sea,

accompanied by warping, so that the later limestones and reefs were lifted

to very different elevations in different parts of the island margin. Since

that event, the whole has been again subjected to erosional attack of the

sea, and to wind work, with the result as now seen in the physical feat-

ures. The comparatively easily destroyed shales, marls and limestones

of the Tertiary series have been extensively removed, leaving only a

fringe of these formations along the north coast and a part of the dis-

tance along the south coast, and developing all of the topographic forms

characteristic of the erosion of emerged coastal deposits, together with

some very special forms due to the peculiar makeup and attitude of the

rocks themselves.

Since this first emergence there have been minor oscillations also, the

record of which is observable in marginal terraces, deeply trenched flood-

plain deposits, and thoroughly indurated wind deposits of presumably

Pleistocene age. Apparently the latest movement has been one of slight

emergence.

A summary, therefore, of the larger items in the geologic history in-

cludes the following: -

1) A long geologic period of voleanic activity, accompanied by marginal at-

tempts at assorting of fragmental and detrital material and organic accumu-

lation disturbed from time to time by renewed or extended igneous activity.

2) A dying out of volcanic energy, greater stability of the mass with respect

to elevation and subsidence, and erosional attack continued long enough to re-

sult in extended planation and partial base leveling with final extensive sub-

mergence.

3) The development -of an unconformable overlying series of shales, reef

limestones and related deposits chiefly of organic origin, brought to an end by

final re-emergence.

4) The development of present surface features under stream erosion and

marine marginal attack, with modifications arising from oscillation of level.

The geologic column forming the basis of this outline, avoiding minor

details that are properly the subject of further study before specific state-

ment should be made, is as follows:

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 61

Recent alluvial deposits.

Submergence with flood-plain deposits.

Younger Series.

San Juan dune sands (Pleistocene).

Submergence with terrace cutting.

‘Post-Arecibo emergence and erosion.

Organic limestones, marls ete. (Mid-Tertiary).

Arecibo reef limestones (Oligocene).

Lares shales ete. (Eocene ?).

Marked unconformity.

Older Series.

Interbedded limestones, tuffs and shales etc. (Cretaceous), Coamo

tuff-limestone, Trujillo Alto limestone, Aibonito conglomerates and shales

with many intrusives.

Interbedded foraminiferal shales and ash shales with tuffs, cut by

many intrusives (Early Cretaceous ?), Fajardo shales, Mayaguez shales,

Barranquitas shaly limestones, Sierra de Cayey tuffs, Comerio consoli-

dated ash beds and tuffs, etc.

FuTURE PROBLEMS

One of the objects of this exploratory study was to discover and define

the problems that should be investigated by this organization. It is not

supposed, in enumerating this list, that these cover every possible subject

of special study, but they do indicate the fields in which there is promise

of immediate and valuable scientific returns, and at the same time will

add to the fund of usable information to be put within reach of the people

of Porto Rico.

BASE MAP

One of the fundamental things as a basis for all sorts of detailed geo-

logic work is a good contour map. The whole island ought to be mapped

in the same manner as is done in the United States, using the same

quadrangle system. On account of the density of population, the com-

plexity of structure and relief and the variety of agricultural uses of the

soil, the scale should be approximately one mile to the inch, or 1: 62500,

so that these maps could be used as base maps for all sorts of special

purposes.

The maps now available are chiefly those of the Interior Department

of Porto Rico, made to illustrate the various reports of the department

and representing the progress of public works such as railways, telegraph

and telephone lines and highways. On account of the care with which

the different classes of roads have been shown, and the general accuracy

of locations, these maps are especially useful in the present investiga-

tions. One of the most useful is a map of the Bureau of Public Works

62 ANNALS NEW YORK -ACADEMY OF SCIENCES

on a scale of approximately 2 inch per mile which has even the kilometer

distances along the roads indicated. Until some sort of a contour map

can be secured. such maps as these will be found eminently serviceable.

Along the south coastal margin within the region of irrigation devel-

opments, there has been some special mapping with contours. In no case

do they cover much ground beyond the outer lowland and terrace border,

and because of this limitation they are not so generally useful for our

purpose as the Interior Department maps. They are, however, very-

much more accurate and detailed and for the territory covered are emi-

nently suitable as base maps.

GEOLOGIC MAP

A geologic map of the island should be one of the results of this series

of studies, whether a relief map is secured or not. Such a map of the

whole island is necessarily an ultimate rather than an immediate product,

but district maps can be undertaken at once, with no difficulty whatever.

These preliminary districts can be selected so as to include some of the

most promising investigation problems in special lines, and both kinds of

work can thus be carried on at the same time. This therefore leads

directly to the next item, which is district studies.

The only geologic map thus far attempied is that by R. T. Hill.

DISTRICT STUDIES

It is possible now to select areas which are known to contain geologic

features of special interest and significance, and it will generally be con-

venient, if not indeed necessary, for the investigator to make a detailed

geologic map as a secure foundation for his special studies. One of these

is the Coamo Springs District, which may be made large enough to ex-

tend from the Descalabrado river on the west. to Salinas on the east, and

reach as far north as Aibonito. It will include as features of special

importance for investigation, in addition to the mapping, the hot springs,

the great conglomerate series, one of the later of the great volcanic vent.

complexes, the genetic history and horizon of the Coamo limestone which

is a striking mixture of volcanic and organic matters, the high floodplam

deposits of the stream valleys and their bearing on late geologic history,

and certain physiographic studies connected with the coastal terraces.

This district promises, as can be seen, an unusually large range of topics

- inviting special study, all of which will be illuminating to further devel-

opment of the geologic survey of the island.

Another district of equal promise in a very different manner is on the

north coast extending from the Quebradillas to the Arecibo river and

ihe

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 63

reaching from the sea to Lares so as to include a strip of the older com-

plex rock series beyond the inner margin of the Tertiary series of reef

limestones and shales which constitute the greater part of the bed rock

of the area. Beside the mapping and detail of structural relations, this

district presents the best opportunity to investigate the question of exact

age of the basal beds of the Tertiary series, the transition from lignitic

shales of perhaps fresh water alluvial origin to massive limestones of reef

type, a subdivision of the Tertiary series, the meaning of the thinning

out and disappearance of the Lares shales toward the east, and a paleon-

tologic study of the beds, all of which are fundamental in any additional

study of other districts containing the Tertiary rocks. There are besides

good opportunities to study the meaning of the high terrace-like shelf

coming abruptly to the sea at Quebradillas river and the meaning of the

deep embayments now occupied by such playas as that at Arecibo. This

is also one of the best localities for a detailed study of the structural and

petrographic features of the San Juan formation as well as the behavior

of modern dune sands along the present coast.

Another district of still different features, and giving foundation for

special studies of quite a different bearing, is that lying between Caguas

and the Caribbean sea and perhaps extending as far eastward as Naguabo.

This will include the largest massive igneous unit in the whole island

and promises information bearing upon magmatic differentiation, origin

of the magnetic iron ores, relation of the great intrusive masses to the

other igneous representatives, petrographic range of the igneous rocks,

and marginal metamorphic or other effects,—studies fundamental in a

final statement of the igneous history of the island.

There are other districts which have special problems associated with

the regular areal geologic work, but these are sufficient to indicate the

range of such district studies and their variety.

Certain special investigations are of a sort requiring comparison and

summary of many different localities, and for these it will not be wise

to handicap the investigator by limiting work to a single district. Some

of these are suggested below. |

REEF-BUILDING ORGANISMS

The limestones of Porto Rico are remarkable for the great prominence

of alge and corals and other closely associated organisms lending them-

selves to the construction of reefs and accompanying deposits. These

forms belong to practically every limestone formation of both the older

and the younger series except those most closely related to the shales. It

is a study requiring the training of a specialist in such lines.

64. ANNALS NEW YORK ACADEMY OF SCIENCES

PALEONTOLOGY

The total organic content is much greater than is intended to be in-

cluded under “Reef-building organisms.” There are immense numbers

of splendidly preserved fossil species of organisms belonging especially

to the Tertiary series. Probably a great many are new to science.

There are probably few places in America or within territory belonging

to the United States where the marine Tertiary succession is of more

promise than in Porto Rico. This problem or line of investigation is

closely related to the next topic, that of Tertiary subdivision.

TERTIARY SUBDIVISION

A faunal and structural summary will naturally lead to the establish-

ing of subdivisions and the determination of horizons in the younger

series of rocks culminating in a statement of the complete Tertiary his-

tory of the island.

SAN JUAN FORMATION

A study of the characteristics and detail of origin and historical steps

associated with the Pleistocene fossil dune sands, referred to as the San

Juan formation, is another problem.

“SUBDIVISION OF PRE-TERTIARY COMPLEX

A discussion of this kind is one that will properly follow upon the

completion of areal work in several of the typical districts. It is, how-

ever, one that will necessitate investigations throughout the interior of

the island, and will include a summary of the characteristics of all of the

prominent local formations. A grouping and correlation cin no doubt

be made in due time.

MINERAL RESOURCES

On account of the interest taken by the people of Porto Rico in the

question of possible mineral resources, it is desirable to undertake an

investigation of the kinds of products, their origin, distribution and prob-

able economic value. Jn connection with this, because of the small

amount of exploratory work that has been done, it would be especially

useful if suggestions were made at the same time about the methods of

exploratory development and the people cautioned concerning wasteful

methods. This work should be done so as to cover the whole range of

mineral possibilities in the island regardless of location. ‘There are

known deposits of copper, iron, gold, lead, silver and zinc among the

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 65

metals as well as a number of non-metallic products. But in no case is

the real value, or the probable extent or the geological relation, known

sufficiently well at the present time to serve as a basis for a discussion.

This should be one of the first undertakings of this survey, both because

of the fact that its value is fully appreciated by the people of Porto Rico

and because its conclusions do not materially depend upon the other in-

vestigations or mapping progress.

PETROGRAPHY

On account of the great number of igneous rock occurrences and the

very great variety that is certain to be shown in quality and minor

petrographic character, and because of the considerable range in compo-

sition already known to characterize the intrusives, there would seem to

be an ample and promising field in this line for a special investigation.

It is possible that some genetic relationship is exhibited in the distribu-

tion of these variations and that a thorough comprehensive study would

throw some light on the more obscure problems connected with the gen-

esis of igneous rocks. This is a problem that can be taken up at any

time, and that need not be regarded as dependent upon special district

studies, although it is evident that the finishing of work on certain dis-

tricts would facilitate a study of this kind.

PHYSIOGRAPHY

Enough is known of the physiographic features and their meaning to

appreciate that a great deal of the detail of the later geologic history of

the island is more or less intimately bound up in the physiographic de-

velopment. The broader or larger physiographic features have already

been suggested, but there are certainly many details, some of which may

well be of much significance in understanding the geologic history, which

will require the special attention of a trained physiographer. Porto Rico

is a unit of geologic history, of geologic structure and of physiographic

form. Each is of sufficient complexity and unity to be made independent

subjects of investigation.

THERMAL WATERS

The hot springs in the vicinity of Coamo Springs suggest from their

situation and reported composition the possibility of being representa-

tives of juvenile waters. It is believed that a study, planned especially

to investigate the origin and character of these waters, together with such

others as may exist, would be a very suitable special investigation. On

66 ANNALS NEW YORK ACADEMY OF SCIENCES

account of the fact that the principal occurrence of this type of water is

very local, it would be possible to combine a study of this kind with a

district study such as has been referred to in a preceding paragraph.

GEOLOGIC HISTORY

The complete geologic history of the Island of Porto Rico cannot be

written until all of these and perhaps other more special investigations

have been made. A complete historical statement must be regarded as

an end product of the whole range of studies carried out for more special

purposes. It is, therefore, the final topic and may well be deferred to, a

time when most of these already suggested have been carried far enough

so that the data of special importance secured by them are available for

this general summary.

It is evident from the appearance of this list that there is a very great

amount of geological work awaiting the investigator in Porto Rico, and

that it is varied and complicated enough to require several years of study

in large part by experts or specialists in all branches of the subject. It

can be seen also that the Island of Porto Rico is a geographic unit of

more than usual complexity and scientific interest and gives promise of

results for effort expended in researches along geologic lines.

COLLECTIONS

A beginning has been made toward securing a representative collection

of typical rocks and fossils. Several hundred specimens were brought to

New York for use in formulating the accompanying description, and as a

possible basis for further more special investigations.

As a first step in this direction, about a hundred thin sections of the

rocks have been made for microscopic study and detailed comparison.

They will form a basis in planning the special petrographic investiga-

tions which may be undertaken.

In lke manner a large number of fossils have been gathered and their

general relations are being studied. Additional investigations along

paleontologic lines will be in large part outlined or suggested by the trend

of these studies, for although the collection is very fragmentary it is

nevertheless characteristic and fairly representative of the principal for-

mations.

- More than a hundred photographs were taken of strictly geological

subjects illustrating typical physiographic features, structural detail of

rock formations, structural relations, etc. These are all suitably labeled

and form the beginning of a collection of ilustrations of Porto: Riean

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 6%

geology. In addition to the regular photographs, a number of photomi-

crographs have been made from the thin sections of typical rocks.

Some of this material will finally serve as a foundation for the geo-

logical section of what it is hoped may become a Natural History Museum

of Porto Rico.

ILLUSTRATIONS

CROSS-SECTIONS

The note books of the party and field maps carry a record of field de-

terminations and detail of structural relations and comments greatly in

excess of what can be published in such a report. They are the property

of the organization and are of particular service as guides in planning

further work and in giving each new investigator his bearings, together

with some suggestions about the character of his own district or the dis-

tribution of data bearing upon his special investigation. The note books

contain observations along some of the principal roads in sufficient detail

to serve as a foundation for complete generalized geologic cross-sections

of the island on two especially important lines.

Cross-sections, therefore, have been drawn to illustrate the kind of sur-

face relief, the grade of the road, the kinds of rocks or rock formations

and the geologic structural relations, and are reproduced to accompany

this report. An immense amount of detail is necessarily omitted or com-

bined into generalizations in order to bring the sections within the scope

of a publication of this kind. It is judged that some of these details will

be suitable illustrative matter for future reports based on studies of spe-

cial districts. One of the sections is based on data gathered along the

road from Ponce to Arecibo. The line is drawn from Arecibo to Ponce

direct and the data are projected to this line. This method tends to

obliterate the windings of the road and secure practically normal propor-

tions and relative positions for the associated formational units. ‘The

other section line is drawn directly from San Juan Point to Santa Isabel.

By projecting to this line all the data gathered on the Bayamon-Comerio-

Barranquitas-Coamo road a great deal more elimination of road curves is

accomplished than in the other section, and it makes the grades of the

road look somewhat abnormal by reason of this shortening of road dis-

tance of certain large curves, but on the whole the relations are shown

without special difficulty except that attendant upon the need of general-

izing the minor structural detail.

68 ANNALS NEW YORK ACADEMY OF SCIENCES

MAPS

A hasty reconnoissance examination is seldom a satisfactory basis for

an areal map. ‘This is quite true of the present investigation.- On this

account, therefore, if it were not for other considerations, an areal map

would not be attempted. But in this case, where a good many more or

less independent special investigations are to be carried on in which a

reasonably accurate geological map will prove decidedly helpful, there is

sufficient excuse for presenting a reconnoissance map. An earlier map

of this kind prepared by R. T. Hill was made under conditions so much

less favorable for travel, and seems to have been constructed in some par-

ticulars with so much less opportunity for observing the actual conditions

in certain areas, that an entirely new map is believed to be the better

solution of the present need. The accompanying reconnoissance map is

intended, therefore, as a convenient guide or location map for subsequent

more special investigations, and it is expected to be wholly replaced by

one of much more detail and greater accuracy as a final product of this

survey.

ACKNOWLEDGMENTS

The members of this expedition have appreciated the very material

help, the sound advice and useful suggestions given by the officials of the

government of Porto Rico, and are indebted to Governor Yeager for his

very practical aid in making arrangements for the field work and for his

live interest in these investigations; to Colonel Shanton, chief of the In-

sular Police, for his willingness to give introductions to men acquainted

with special mineral localities and for his precautions to insure protec-

tion against unnecessary delays; to Mr. Wheeler, of the Interior Depart-

ment, for assistance in securing suitable maps as a basis for travel and

notes; to Mr. Bonner, the Auditor, for facilitating the settlement of ac-

counts; to Dr. Lippitt, of the Bureau of Sanitation, for information

regarding sanitary precautions and hotel accommodations; and to Mr.

Campbell, of the Bureau of Transportation, for the excellent equipment

for travel which contributed largely to the success of the expedition.

Many others have been of assistance in pointing out localities of special

interest, in giving names of reliable informants and in acquainting us

with the usages and customs of the country.

BERKEY, GEOLOGICAL RECONNOISSANCE OF PORTO RICO 69

BIBLIOGRAPHY

The following papers contain descriptions of physical conditions and fea-

tures and of some geologic observations in Porto Rico. A suggestion as to the

range or character of the article in most cases follows the title.

CLevE, P. T.: “Geology of the Northern West Indies,’ with maps. Kongl.

Sevenska. Vetenskaps, Akad. Handlingar, vol. 9, No. 12. 1871.

: “Outline of the Geology of the Northeastern West India Islands.”

Ann. N. Y. Acad. Sci., vol. 2, pp. 185-192. 1883.

Davis, Bric. GEN. GEORGE W.: Porto Rico—embracing reports of Brig. Gen.

George W. Davis, arranged by topics. Government Printing Office, Wash-

ington, 94 pp. 1900.

Dinwippiz, W.: “Physical features of the Island” (Puerto Rico). MHarper’s

Weekly, vol. 43, p. 248. Mar. 11, 1899.

: “The great caves of Puerto Rico.” MHarper’s Weekly, vol. 48, p. 293.

Mar. 25, 1899.

DoMENECH, M. K.: “Mineral resources of Porto Rico.” Mines and Minerals,

vol. 19, pp. 529-532. 1899.

FALCONER, J. D.: “Evolution of the Antilles.” Scot. Geog. Mag., vol. 18, pp.

369-376, 1 pl. 1902. Discusses general geologic history of America, and

especially that of Central America and West Indies.

Guppy, R. J. L.: “Geological Connections of the Caribbean Region.” Trans.

Canadian Institute, pp. 373-391. January, 1909. Discusses general re-

gional relations and the probability of an Antillean continent.

Fiske, Amos K.: “The West Indies.” G. P. Putnam’s Son, New York. 1899.

Folded maps, plates. Physical characteristics of Porto Rican history ete.,

social and economic conditions.

HAMILTON, S. HARBERT: “Notes on Some of the Ore Deposits of Porto Rico.”

Eng. and Min. Journal, vol. 88, p. 518. September, 1909.

Hitz, Ropert T.: “Cuba and Porto Rico with the Other Islands of the West

Indies.’ New York. 1898. A book of over 400 pages containing a very

large amount of information covering many matters in addition to the

physical features and the geology.

: “The Value of Porto Rico.” Forum, vol. 27, pp. 414-419. June, 1899.

: “Porto Rico.” National Geographic Magazine, vol. 10, pp. 93-112.

1899. One of the best articles on the physical features of Porto Rico.

: “The Geology and Physical Geography of Jamaica.” Bull. Museum

Comp. Zool., vol. 34. 1899. <A study of a type of Antillean development.

: “The Forest Conditions of Porto Rico.” Bulletin 25, U. S. Department

of Agriculture. 1899. Quoted in part in Public Opinion. Contains very

brief description of physical features and has a good relief map.

NEWBERRY, J. S.: “Geology of the West Indies.” Trans. N. Y. Acad. Sci., vol. 1,

pp. 23-24. 1882.

Nitze, H. C. B.: “Investigations of Some of the Mineral Resources of Porto

Rico.” Twentieth Ann. Rept., U. S. Geological Survey, Part 6, pp. 779-878.

Gives a tabulation of the mineral occurrences, and a very brief note on

the principal ones.

70 ANNALS NEW YORK ACADEMY OF SCIENCES

SPENCER, J. W.: “On the geological relationship of the volcanics of the West

Indies.” Victoria Inst. Jour. Trans., vol. 35, pp. 189-207, 1 fig. 1903. Dis-

cusses physiographic features and changes of the West Indies and the

submerged platform upon which they rest, place of their igneous forma-

tions in geological history, ete.

VAUGHAN, T. W.: “Earliest Tertiary Coral Reefs in Antilles and United States.”

(Abstract.) Read before Geol. Soc. Wash. Feb. 26, 1902. Touches on

coral reefs of Porto Rico being in part of Oligocene age. No Miocene

reefs, because in Miocene Antilles stood at higher level than now. Plio-

cene reefs in Florida and Antilles.

WuHarton, W. J. L.: “Depth of the Ocean near Porto Rico.” Geog. Jour., vol.

4, p. 255. Sept., 1894. The physical condition of the ocean.

WILSON, HERBERT M.: “Water resources in Porto Rice.” Water supply paper

No. 32, U. S. Geol. Surv. 1899. An excellent discussion of the physical

features. Contains many good illustrations and two maps with reconneis-

sance contours.

: “Vhe Engineering Development of Porto Rico.’ Eng. Mag., vol. 17,

pp. 602-621. July, 1899.

: “Porto Rico, its Topography and Aspects.” (With map.) Bull. No.

32, Amer. Geog. Soc., pp. 220-238. 1900. A good description of the physical

features, climate and productive industries.

Author unknown: “Porto Rico, its Natural History and Products.” Scien.

Am. Suppl., vol. 46. July 30, 1898. No author mentioned—good general

account, including report on geology and mines.

Author unknown: “Zur Kenntniss der Insel Puerto Rico.” Geographische

Gesells. Ham. Mett, 1899-2, pp. 217-236.

Author unknown: “North American and West India Gazetteer.’ London,

printed for G. Robinson, 1778 (6) xxiv, 216 pp. Situation, climate, soil,

produce trade—bays, rivers, lakes, mountains, number of inhabitants, ete.

Maps. 2d edition.

ANNALS N, Y. Acap. Sci.

CEOLOGICAL RECONNAISSANCE MAP

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American Museum of Natural History, —

ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

“Vol. XXVI. pp. 71-148

Editor, Epmunp Otis Hovey

LIST OF GREATER ANTILLEAN SPIDERS

WITH NOTES ON THEIR

DISTRIBUTION

@. BY

ae i Frank EK. Lutz

: }

ee S , NEW YORK

_ — + PUBLISHED BY THE ACADEMY

eo a 29 May, 1915

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Natura History, 1817-1876)

OFFICERS, 1915

President—GerorGE FREDERICK KUNZ, 601 West 110th Street

Vice-Presidents—CHARLES P. BerKEY, RayMonpD C. OsBURN,

| - CHARLES BASKERVILLE, CLARK WISSLER

Corresponding S ecretary—HENRY E. Crampton, American Museum

Recording Secretary—EDMUND Ors Hovey, American Museum

Treasurer—HENRY J. CocniRan, 389 Fifth Avenue

Librarian—Ratru W. Tower, American Museum

Editor—EpMunpD Otis Hovey, American Museum

- SECTION OF GEOLOGY AND MINERALOGY

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ERRATA ©

ANNALS N. Y. Acapb. Scr., Vol. XXVI

Page 144, first column, line 20 from top, for coronotus read coronatus

** 144, second column, line 18 from top, for Ellica read Hilica

145, first column, line 9 from top, for 60 read 101

147, second column. line 10 from top, for Salticus read salticus

a ¥ 2 ; ae “2 wae os

pon Bet 4 UM

; we cy, |

ah PON A, eG ia fee.

ere el) Baas ah

[ANNALS N. Y. AcapD. Scr., Vol. X XVI, pp. 71-148. 29 May, 1915]

LIST OF GREATER ANTILLEAN SPIDERS WITH NOTES ON

THEIR DISTRIBUTION }

By FrRanK E. Lutz

(Presented by title before the Academy. 12 April, 1915)

CONTENTS

Page

Eb PEERS IES Se eS Oe On eee Seg oe, a ae a a a aire a (2

et SE LEO So ee ee oo ee 75

SNOPES Seis koe tele Cae fo sata fae A ate hs wis Gh aunrce we eae Siam a weaig ome 75

EPRINTS Pe eae sft a Pe St PPI Be SAO ie ORIG mamas Oe OR eae 75

ae RE ea Ey Re, Toe se bie SATS ae Re OS earl 76

RMMNMNCDD NE DIME SENET oh ei en ere oR: eve hss Sia, Lg isa Ria a waake Dans erate 76

LTE STILTS NB ee ie EE ORS a Perea Se ee nee Re a Por A gan rir

Aranes Verz............ ON STS ny op og Po eT ae er ea ae oe ee ee ae ria

| SET EEL Sie pa i NS ae oe Aine NE RS A ge eal Soe er 78

SERED RR ne cre at ee hte CS eS wihace SAT, mies) Uae See Nes 78

Oe) LERENT SU fame |e Acai ee ce VR ae ae eS RANE DN Te fe DISH rv Oe 78

ie SECURES SERRE ERENCE er Acris Nye Gag eae steak Gu) a wa dae oe Bee hs ee 79

MRE ee at tes is a ares ES eS Peck cya ow a Win Daley aan reine 79

ENN SE eee eet ies ne ne eee Meola kc Sede bain oe We mae oe baat aide 79

NaN NSUNU BAe eit fens SS A OIL 2s oon dl wae! Gia, 6 alba Sw ae ee me Soe eg SO

NII EPI gate Seta eS a oe ES oo tas ob id eh A ate age eee SO

ue NROM AIUECE Te rates nce ee Shaye ho Ee aera he SI oid oS a va Si Seach Dein dim ake RE SLL 81

02 TE PUD EEE sD ual Te ee ae SR eC a on eR og Oy 81

NE em gr a) Scent 9 ae Sen Se SS os Se awa ew ertiaa atk aS Siehetans 81

RT as pi ee Chara Sorcha we Se ha had Oe OS bE ae wee ee 81

Drassidz...... OF RN oa pe AB MS gy nT ee 4s Oe Bre ee akin Paks Bite 2

0] LELETEE ES S8 Satis AR 5 eee dace i Oe Date at nec pai eee Cane ee aR 82

LL LL ED LSE SE 2 Se ee One oe ae ae oe ae aa ee et en ne SS ety 82

er NNN Ta ec Wet arr ES cee ay ae Soe A a Ale tes Saha wi Eevee MEMES Ree We ROR s+

RNR oe a ee eee s pickle Ce mc wis Couns FG BO Met Meee Ee 86

SMES ret: IED St aN e O ote hte ed Sethe ort s Sh cin cia vaveie ave od Ols yma tne ete eiele ri

2) SSUES Sy LTT eS eae ee Cn eR eT ee eee Se ee eee er eee 2 88

eee SOPOE MANNE Seb oct pee tc sat ckaen so hos Siete Wika Ga! idly wcantaa sinks amare buass bane 89

2 ES CET eee ae EC ©, See ee ee eR Men nCeE ae te Abe neler 90

NEE NER ai ee and Ne Le ee New Rers, Gs etalk al eh 7e, lee EM ate ere ia ei trea we 95

REI RMS ns Po Sh) dams ai ome ea rene aS eh es te RL ie Pa oe ae 95

MRE AEAEICTUNERS 5 250 IP PA Kalan phe hoecctiee ake os Sik es ahaw Bi ato WEIR we Ie 95

SaaS UE MESSE FISD ttc Se oe ets lass bine ata Baie (elev Soe § Pei Semis Cas 96

SST ST SST 025 2 ae ee Bo i OD pe SN SUA ace ie A Oe oe Bee ae ce 96

UURMUREERR SES he ethers eae Pel ie te Mes Loe wig Oe ork eal ees Rng ok a a 97

1 Manuscript received by the Editor, 1 April, 1915. Ps

79 ANNALS NEW YORK ACADEMY OF SCIENCES

Page

Selenopines foc es a ae oe ee ee. 97

Sparassinvey 2.25354 Ce ee Eee ee ene & a. 97

Clibioni nee: 2.02 2) oo ee eee Be ee), 98

Chenin 555 3S a SRS ea. Se eas ee ee ee eee 99

Lideraninwi... os E28 oe ete eee eee Sra eo ee ee 99

Micariima.. 6-255 id eee eee ae ee ce on 99

Corinming . 66.5025 be oe FS ee ee Wee eee oc ee. 100

Avelenidee. o.oo) 22 anc ee a oe ee eee ee eee 100

Pissarides § og.sspl eile .o Oe Fee ala ere ee es ee 101

LY COSTAR. 3.6555 5 ES ss soe ae OS ee ee ee ee ee 304:

OXYVOPIG ose elk oe eee ne bre Re ee ee 102

Salers) A. Soe oer, oe es ee ee ee ee (nid Sooke ee 103

Distribution of families... ..24 2. suds ous acme ee oe Oe eee 108

Distribution of genera. : .. 056. nse os Sea bs cles ce oe eee ee 110

Lesser Antilles. .9o.c.. ete ee el pete es eae 6 oe oe 110

POFrto Ri@0s.c2. 3 ee eee eee Pe Oe RIES aOR Fe a an PS ie Oe 4453

FRI aril. ic eo 5 8S bow, on ic wedi ane Siw ale es eee ee 115

GODS 2 oss cos 6 6 ae ee Sie lew ete so ape Stee leas cnn 116

oF AYMAICH 5 op. ae on Swis oad 0 01m wrsoe es eee w ele eee One eRe 118

Greater Antilles... oss 40.0. 32 bee Cee Soke Se eee 120

Distribution ‘of specresis 3 ke ee eee oe Bite hee 122

Genéral ‘discussion..c 286.50 oF 30s soak oe eee "« wire eh 422

Is the fauna of the Lesser Antilles distinct from that of the Greater

ADITIMOS ? 5 a. oicdeck wee Soe 5 Sie wok. cms Sei ee ee ime cee £22

Is the Antillean fauna distinct from that of the mainland?.......... 124

Mainland affinities of the Antillean fauna....:................eeeeee 27

Origin of the Antillean falina...t.. 2225... s¢..0-5 = 2) 3) 133

SUMNINALY 6 5. ses oe 5d ois ew a Bw bye tare Ra eee 141

Bipliography®. ... <6. 6s as ae « & te ews le ere as oer ee 142

PING OR oo foc alee wie oo bin ate Haledon eters nee ee Ae 148

INTRODUCTION

The principal general lists of Greater Antillean spiders-are by Nathan

Banks, who has reported on collections from Cuba (1909), Haiti (1903)

and Porto Rico (1902). Practically the only list for the Lesser Antilles

is Kugéne Simon’s for St. Vincent. Except for Banks’s Cuban list, which

was overlooked, A. Petrunkevitch (1911) has included nearly all the

published records. For the sake of uniformity J have followed rather

closely his synonymy even though it is not, in all cases, in agreement

with the ideas of Mr. Banks, who is responsible for the identification of

most of our specimens. Unless otherwise stated, I have also relied on

Petrunkevitch’s catalogue for data concerning distribution in the western

hemisphere. I have usually departed from the alphabetical arrangement

of genera used by Petrunkevitch and followed more nearly the arrange-

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS "3

ment of Simon’s Histoire Naturelle (1892-1903) and have rehed largely

upon Simon for data concerning distribution in the Old World.

The dissertations on distribution have been included because a study

of the problems of distribution is the main excuse for a faunal list. I

well realize that we know far too little of the fauna of this and neighbor-

ing regions to attempt to draw final conclusions and I also realize that

even if our knowledge of the spiders were perfect it would be necessary

x *}

Fic. 1.—View of a portion of Mona

Nearly all of the specimens referred to in this paper came either from the grassy

portion of the coastal strip (shown in the foreground) or from the thicket at the base

of the cliff,

to study, in connection with their distribution, the distribution of other

organisms, especially those of which we have more geologic records, but

it seems that we have now a basis for an interesting discussion at least.

‘The new records in this list come mainly from four sources, the speci-

mens being in the American Museum of Natural History: collecting in

Cuba in September and early October, 1913, by the author and Mr. C. W.

Leng; in Porto Rico, Mona (Figure 1) and Desecheo (Figure 2) in Feb-

ruary and March, 1914, by the author; in Jamaica in February and

v4 ANNALS NEW YORK ACADEMY OF SCIENCES

March, 1912, by the late Mr. J. A. Grossbeck: and in Culebra and Porto

Rico by Professor Wm. M. Wheeler. The first two collections were iden-

tified by Mr. Nathan Banks. Such of the last two as have been worked

up were largely identified by Dr. Alexander Petrunkevitch. Although

all of these expeditions were primarily for insects, careful field notes ac-

company most of the spiders, and these, when deemed important, have

been incorporated here.

Fic. 2.—View of a portion of Desecheo

Unlike Mona, Desecheo has no coastal strip. The cliffs. however, are not as precipitous

as on Mona. They are, for the most part, covered with trees or a xerophytie thicket.

Where a more or less definite tvpe locality for a species is known, it

usually has been indicated by the sign + before the locality, unless it is

given by the context. As far as possible, the entire range is given in

every case, so that when only one locality is mentioned it means that the

author knows of no other records, but frequently an inclusive term is

used, such as “United States,” when a number of more definite localities

are known.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS "

ARANEA THERAPHOSA

Liphistiide is an interesting family which is sometimes and on good

erounds given the rank of a suborder (Mesothele). It has but two

recent species, both of which belong to a single genus (Liphistius) and

are known only from the East Indies. The fact that they have segmented

abdomens and that they were apparently the predominant group in

America and elsewhere in Paleozoic times indicate a great antiquity.

If it be true, as I believe it to be, that the West Indies share with

Oceanica, Madagascar and parts of Africa and South America the honor

of being preserves for ancient Aranez, it is possible that representatives

of this group may yet be found in the Antilles.

AVICULARID®

This family is variously known scientifically. Unfortunately the pop-

ular name for certain of these creatures is tarantula—a term used in

scientific literature for a genus of whip-scorpions occurring in the West

Indies and elsewhere. It is curious that no Paleozoic remains of this

family have been found in either North America or Europe. The large

size of the recent forms and the undoubted age of the group would lead

one to expect such fossils. Of the seven subfamilies, one (Miginz) is

confined to Africa, Madagascar and New Zealand: one (Paratropidine)

is found in the Amazon region and in St. Vincent; one (Actinopodinz)

has one genus with a unique species in southern Africa, one genus in

Australia and one in Central and South America. The other subfamilies

have representatives in the Greater Antilles and will be considered more

in detail.

Mygale is a preoccupied name but there were a number of species

described under it which have not yet been placed elsewhere. Of these,

maculata Walckener is from Jamaica and the following were described

from the West Indies: athletica Koch, probably an Hurypelma; conformis

Koch, a Trechona, according to Ausserer.

CTENIZINA

These are the “trap-door spiders,” so called because many of them

close the entrance to their underground tunnels with a hinged lid.

Pachylomerus is found not only in America from the District of

Columbia to Venezuela and possibly Brazil, but also in the Mediterranean

region and Japan. One species is known only from St. Vincent and the

only other Antillean species, so far as is known, is nidulans (Fabricius).

It is probably confined to Jamaica where it is rather common in the

76 ANNALS NEW YORK ACADEMY OF SCIENCES

central parts of the island in places where the “soil is not exposed to the

effects of the extremes of heat and moisture” (Sells).

BARYCHELINZA

Stothis. Petrunkevitch lists two species from Venezuela and one

from St. Vincent but he overlooked cubana Banks from +Santiago de las

Vegas, Cuba.

Trichopelma, has two species in Brazil and one, nitidum Simon, con-

fined to Santo Domingo. |

.The two genera just mentioned belong to a section of the Barycheline

which are tropical American except for a genus from the Mediterranean

region and eastern Africa.

AVICULARIINA

Ischnocolus has, according to Petrunkevitch, no representative in

America. JI. hirsutus Ausserer, recorded from +Cuba and the Bahamas,

is believed by him to be a young Lasiodora. Banks records the genus

from 2,800 feet altitude on El Yunque, Porto Rico.

Stichoplastus has one species recorded from each of the following:

Venezuela, Trinidad and Guatemala. Professor Wheeler took a speci-

men, as yet unidentified, in Culebra Island, Porto Rico.

Hapalopinus has only one species, cuwbanus Simon, and it is confined

to Cuba.

Scopelobates sericeus Simon is the only species of its genus and is

known only from its type locality, Puerto-Plata, Haiti.

Cyrtopholis has one species in Mexico, two in South America, one in

the Bahamas, and three in the northern Lesser Antilles. In addition,

agilis Pocock and cursor (Ausserer) have been recorded from Santo

Domingo; innocuus (Ausserer) from +Havana, Cuba; and jamaicola

Strand from Montego Bay, Jamaica. It belongs to a group which is

entirely American, as far as is known.

Eurypelma. To this American genus belong the large tarantulas—

the creatures which are usually thought of when the latter name is used.

Only one species has been recorded from the West Indies. It is spinicrus

(Latreille) from Cuba and is fairly common there. Banks states that

he has seen an immature Hurypelma from Cayamas, Cuba, which repre-

sents another species but was unable to place it. The other thirty species

are found chiefly from southwestern United States to northwestern South

America.

Avicularia is largely confined to South America, there being one

species reported as confined to California, one to Panama, three to the

LUTZ, LiST OF GREATER ANTILLEAN SPIDERS V7

Antilles and fourteen to South America. Of the Antillean species, one

is recorded only from Guadeloupe and Martinique while two, c@sia

(Koch) and l@ta (Koch), are known only from Porto Rico. Banks

recorded /wta from Culebra and Utuado,? Porto Rico.

Phormictopus cancerides (Latreille) is the type of a genus the only

other known species of which is probably found in South America. P.

cancerides is reported from Brazil, tSanto Domingo, Culebra,* and the

following Porto Rican localities: Lares, Anasas, San Juan, Hacienda

Esparanza and Isolina.

DIPLURINZ

Diplura macrura (C. Koch) is spoken of by Simon as confined to

+St. John. Banks records it from Pinar del Rio, Cuba, under the name

of Ischnothele. ‘The remaining members of the genus are from South

America and three closely related genera are confined to that continent.

Two other genera are found in South America and also in Madagascar

and Australia, one of them reaching even parts of Asia and Africa.

Ischnothele digitata (Cambridge) is recorded from Mexico and +@Gua-

temala. It is said to live “in holes in the ground, near which it makes

small, strong, irregular webs, consisting of various floors or chambers.”

Mr. Grossbeck took it near Cinchona, Jamaica, under-a rotten log. In

America the genus is reported from Mexico to Bolivia and also in the

Bahamas. Either it or a closely related genus, T’helechoris, is found also

im Madagascar, East Africa and India. Of the group of genera to which

it belongs, another is American, two are confined to Australia, one to

New Zealand, one to New Caledonia, one to the Trans-Caspian region,

and one is found in Spain, Malasia and New Zealand.

Although Accola is not known in the Greater Antilles, its distribution

is worth mentioning, namely, St. Vincent, Venezuela fae the Philippines.

A closely related genus is confined to the Fiji Islands.

Atypide is a small family which ranges over much of the warmer

portions of the northern hemisphere and is even found in Argentine and

Malasia. It is not known from the Antilles.

ARANEAG VERA

The Hypochilide are undoubtedly Aranee Vere but they differ from

other members of this suborder and agree with the Theraphose in the

‘2The spelling throughout this paper (Proc. U. 8. Nat. Museum, XXIV, 1902) is

“Utado."’ Doubtless the place now known as Utuado is meant.

3This and the Porto Rican records are given by Banks under the name of Schizopelma

erichsonii (atreille).

vas ANNALS NEW YORK ACADEMY OF SCIENCES

possession of two pairs of book-lungs. There are but three species ac-

cording to Comstock, one in the mountains of Tennessee, one in China

and one in Tasmania.

ULOBORID

Like the Argiopidz, the members of this family spin orb webs but the

webs contain a hackled band not found in those of the Argiopids. The

spiders themselves differ in important anatomical characters. Three of

the five American genera occur in the Greater Antilles.

DINOPINZA

Dinopis is found in Africa, Madagascar, Australia and certain of the

Pacific Islands. Of the eleven American species, two are recorded only

from Mexico and Central America; seven from South America; and one,

spinosa, from southeastern United States, Venezuela and St. Vincent.

D. lam1a Macleay is recorded from +tCuba. The only definite localities

given in this island are Santiago de las Vegas (sweeping grassland) and

Cayamas. We took it on the steep rocky coast of Desecheo by beating

low shrubs. Dinopis belongs to a subfamily of which the only other

genus is found in Africa, Australia and New Caledonia. It is evidently

an old group.

ULOBORINE

Uloborus is cosmopolitan and U. geniculatus (Olivier) is known from

Australia, Malay Archipelago, Bourbon, Bermuda and the American

tropics and subtropics. This species is recorded from several of the

Lesser Antilles but the only definite records which have come to my

attention from the Greater Antilles are the following by Banks: the

laboratory at Santiago de las Vegas, Cuba; near Port au Prince, Haiti;

and Lares, Porto Rico. It is surprising that we have not taken it in

the Greater Antilles. It is usually found about houses and its webs are

frequently conspicuous by reason of star-like egg-sacs fastened to them.

U. americanus Walckener is widely distributed in the western hemisphere

but is more northern in its distribution than geniculatus. We have it

from Labrador and it is known throughout the United States and south

to Costa Rica; also in the Bahamas. The only definite Antillean record

I have seen is that by Banks, under the name of plumipes, from Cayamas,

Cuba. U. republicanus Simon is known from + Venezuela and Cuba. We

took it at the edge of a mangrove swamp near Cabanas, Cuba, and ob-

served the same habit noted by Mr. Schwartz (1904) at Cayamas, Cuba.

He says: “Each spider has an individual web, but all are placed in a

great communal web, one of which was 7 to 9 feet wide, 5 to 7 feet high,

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS V9

and 3 feet in depth. The male spiders were in one of the lower corners

of this common web.” There were about 1,000 spiders in this web and

several other smaller ones had about 300 spiders in each. Of the re-

maining American species of Uloborus, one is confined to California,

nine to Mexico and Central America and fourteen to South America.

MIAGRAMMOPINZA

Miagrammopes is known from Africa, Madagascar, southern Asia

and Australia. It has two species in Mexico, one in Guatemala and four

in northern South America. Banks recorded the genus from Haiti on

the basis of an immature specimen which could not be placed in a given

species. He also described cubanus from Cayamas, Cuba. The remain-

ing species 1s scoparius Simon, which is recorded only from St. Vincent

but which we took near Arecibo, Porto Rico, and on Desecheo. In both

these places it was among shrubs and low trees. This genus belongs to

a subfamily which contains but one other genus (//yptiotes) and it is

northern in its distribution, being known only in Europe and America

north of Mexico except, possibly, for one species from Ceylon.

DICTYNIDZ

This family is represented by about a hundred species on the American

mainland but seems not to have been reported from the West Indies.

We took an undetermined species of Dictyna at Guane, Cuba. This

genus is found in Europe and the Mediterranean region, northern and

central Asia, Japan, the Philippines and from the extreme northern to

the extreme southern parts of America. In America it is rather northern

in its distribution, twenty-two species being known north of Mexico as

compared with thirteen in South America.

CECOBIID =

This family contains but one genus, Gicobius. It is known from the

Azores, Canaries, the Mediterranean region, Arabia, Japan, New Cale-

donia and America. Three of the five American species are apparently

confined to South America; one to Florida; and the fifth, partetalis

(Hentz), is recorded by Petrunkevitch from Massachusetts, Florida,

+Alabama and Lower California. Simon makes this species a synonym

of navus Bl. and records it from the Atlantic islands, Japan, New Cale-

donia, Venezuela, southern United States and the Antilles. I do not

know the basis of the last locality. He says it is undoubtedly carried

by commerce. We took it under the loose bark of a stump on the coastal

80 ANNALS NEW YORK ACADEMY OF SCIENCES

flats of Mona. In the United States it lives in crevices of walls, espe-

cially those of buildings.

FILISTATIDE

This family also has but a single genus, Filistata. Jt is found im

the Atlantic islands, Mediterranean region, Africa, central Asia, Philip-

pines and Australia. In America one species is reported from the

Galapagos Islands, one in Peru, one in Guatemala, one in Mexico, and a

fifth. hibernalis Hentz, from t+southern United States south to Brazil,

Paraguay and Argentina. This species is known also from Bermuda;

Jamaica; Havana. Santiago de las Vegas, and Cayamas, Cuba; Isle of

Pines: San Juan, Porto Rico: and St. Vincent. We took it at Banos San

Vincente, Cuba. It lives about houses and under stones.*

The families of true spiders thus far considered are provided with

cribellum and calamistrum. Those which follow lack these structures.

SIcARIID®

This family is also called Scytodide. The six genera are so distinct

that Simon considers each to be a separate subfamily.

Drymusa has but two species. One is known only from Cape of Good

Hope and the other, nubila Simon is recorded only from St. Vincent.

We found it fairly abundant under bits of wood and stone on Mona.

_ Scytodes is distributed throughout most of the world including Mada-

gascar and Polynesia but apparently not Australia. Some of the species,

normally living under stones and logs, have taken to living about build-

ings and have probably been transported by man. S. bajula Simon is

recorded from Mexico; + Venezuela; Havana, Cuba; and St. Vincent. 8S.

fusca Walckener has been reported from Bermuda; Mexico to +Guiana

and Brazil; Havana and Santiago de las Vegas, Cuba; Haiti; a cave

near Pueblo Viejo, Porto Rico; and St. Vincent. We found it under

fallen leaves in a mangrove swamp near Cabanas, Cuba, and under bits

of wood on the coastal plains of Mona. S. lineatipes Taczanowski is

known from Mexico, +northern South America and St. Vincent. We took

it in weedy vacant lots at Santiago, Cuba, where it had probably been

brought by commerce. S. longipes Lucas is found on Bermuda; Baha-

mas; +Mexico to Brazil and Paraguay; Havana and Santiago de las

*Since writing the above I have heard from Mr. Banks that he has seen F. insignis

Cambridge from Santiago de Cuba. Cuba. This is the species referred to above as being:

confined to Guatemala. It is not included in the discussion at the end of this paper.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 81

Vegas, Cuba; Haiti; Lares and Luquillo, Porto Rico; and St. Vincent.

We took it in a hotel at Mayaguez, Porto Rico.

LEPTONETID#.

Two species of Ochyrocera and one of Theotima have been found on

St. Vincent but no member of this family is reported from the Greater

Antilles. It is worth noting that there are only two species in Theolima.

One is confined to caves in the Philippines (liuzon) and the other is

found under vegetable debris in St. Vincent and Venezuela.

OONOPID-E

This rather large family is represented on St. Vincent by fourteen

species, nine genera, but there are no records from the Greater Antilles.

It is found throughout most of the tropical world and invades the tem-

perate regions to some extent. We took a number of unidentified

Oonopids at Banos San Vincente, Cuba, by sifting leaves from the base

of the cliffs. Sifting fallen leaves on the steep sides of Desecheo turned

up unidentified species of Dysderina and Opopza and the same method

of collecting found Dysderina sp. in a sea-grape thicket at Dorodo, near

San Juan, Porto Rico. Both of these genera have been found in St.

Vincent, as well as in the warmer parts of the mainland, Africa and Asia,

including the Philippines.

DYSDERID.®

This family has four small genera in America of which Ariadna is

the largest and the only one known from the West Indies. This genus

is recorded from the Mediterranean region, southern Africa, Japan,

Sumatra, Australia, Tasmania, Massachusetts to Uruguay including Gala-

pagos Islands, and one species from St. Vincent. A. bicolor (Hentz),

which is found in eastern United States from Massachusetts to Alabama,

was taken by us at Banos San Vincente, Cuba. Although this species is

largely a “house spider” and, so, liable to accidental dispersal by man,

the fact that this Cuban locality is in the interior and not along ordinary

lines of travel makes it seem probable that this species is well established

in Cuba. A. solitaria Simon is the species which has been recorded from

+St. Vincent. We found it under fallen leaves in a sea-grape thicket on

Desecheo.

CAPONIIDE

There are only three genera in this family. One genus is represented

by a single species in South Africa. The other two are exclusively

2 ANNALS NEW YORK ACADEMY OF SCIENCES

oa)

American. Caponina has one species in Guatemala, one in Venezuela

and one in St. Vincent. Nops has twelve species of which two are con-

fined to Lower California, one is in Central America and Colombia, and

seven are known only from South America. VV. guanabacoe Macleay

is known from Central America, and +Guanabacoa and Santiago de las

Vegas-in Cuba. V. coccinea Simon is recorded from Haiti and +St.

Vincent.

DRassID£

Members of this large family usually live under stones, bark, or in

other crevices. The American species are, for the most part, northern.

It has been split up into subfamilies and these into groups, but it does

not seem desirable to note these here. It should, however, be said that

Teminius insularis Keys. which has been considered a Drassid, is now

called Syrtsca keyserlingt Simon and classed among the Clubionids.

Callilepis is found throughout much of the Old World. There is one

species found both in Europe and Canada, two on our Pacific Coast, one

trom New Hampshire to Florida, and Banks has recently described grisea

from a specimen which Mr. Leng beat off a pine tree on the sandy plem

south of Pinar del Rio. Cuba.

Eilica cincta Banks is known only by the original material from-

Havana, Cuba. Of the other two species of this genus, one is ieund i

Florida and one in Brazil.

Sergiolus has four species reported only from Canada and the United

States and one only from St. Vincent. We took an immature specimen

south of Pinar del Rio, Cuba. As rariegatus (Hentz) is fairly wide-

ranging and occurs in Florida, this may be the species. It should be

said that the sandy plain south of Pifar del Rio reminds one very

strongly of Florida.

PALPIMANID

Of the four American genera, three are confined to South America

and are each represented by a single species. Otiothops has two species

in Venezuela, one in St. Vincent and walckeneri Macleay in Cuba.

We took it under fallen leaves along the base of cliffs at Baiios San Vin-

cente, Cuba. This genus belongs to the subfamily Palpimaninz, whose

range in the Old World is Africa and southern Asia.

PHOLCID#

These long-legged spiders spin irregular webs, usually in dark places.

They are, for the most part, southern in their distribution. All, except

an Arabian genus, belong to the subfamily Pholcinz.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 83

Artema is a genus well separated from the rest of the subfamily and

contains a small number of widely distributed, closely related species. It

is found throughout Africa, including Madagascar, tropical Asia, Mala-

sia, Polynesia, and tropical America. A. atlanta Walckener is the only

American species. It is recorded from Mexico to +Brazil and Paraguay ;

St. Vincent; and Utuado, Porto Rico.

A group of six genera are separated from the rest of the subfamily by

Simon under the name Blechroscelee. _They are all confined to tropical

America except one which is found as far north as the District of Colum-

bia and also in India and the Pacific Islands. Two of these genera,

Blechroscelis and Mecolesthus, are found in the Antilles.

Blechroscelis serripes Simon is recorded by him from the West Indies

in his Histoire Naturelle but no definite locality is given. It is probably

a nomina nuda. The other species of the genus are in Brazil and

Colombia.

Mecolesthus signatus Banks was described from specimens taken

along a mountain stream in the foothills of tH] Duque, Porto Rico. An

undetermined specimen of the same genus was found on the moist cliffs

along the road from Arecibo to Utuado, Porto Rico. The only other

species in the genus is known only from Venezuela.

Modisimus and Hedypsilus form another group. Both are tropical

American. J/odisimus is known from Texas to Costa Rica and

(glaucus Simon) from St. Vincent, St. Thomas, Santo Domingo and

Jamaica.” No definite type locality is given.

Physocyclus globosus (Taczanowski) is noted by Banks from Santi-

ago de las Vegas, Cuba. We took it in the same island about plantation

buildings at Cabanas and Guantanamo. It has been recorded from south-

western United States, Mexico, Colombia, ¢Guiana and St. Vincent. The

genus is known also from tropical Asia and Africa.

Smeringopus elongatus (Vinson) is found in the tropics of both

hemispheres but the only definite records from the Greater Antilles which

have come to my attention are Santiago de Jas Vegas, Cuba; and Haiti.

Both are by Banks under the name of Pholcus tipuloides Koch. It

appears that no true Pholcus, a genus belonging to a different section of

the subfamily, has been reported from the Greater Antilles, but the cos-

mopolitan P. phalangioides (Fuesslin) will undoubtedly be found.

Smeringopus is a widespread tropical genus.

>Since writing the above I have heard from Mr. Banks that he has seen it from

Havana, Cuba. It is not included in the discussion at the end of this paper.

4 ANNALS NEW YORK ACADEMY OF SCIENCES

‘THERIDIIDE

The members of this large family spin irregular webs and certain of

them are very common about human habitations.

Argyrodes is found throughout the tropics and subtropics. We took

what Mr. Banks identified as A. /arvatus Keyserling in a narrow steep

ravine near Banos San Vincente, Cuba, and an unidentified species, prob-

ably the same, in a similar ravine near Merciditas, a few kilometers far-

ther north. This species is now considered to be a synonym of cancellatus

(Hentz) and the range is Connecticut to +Alabama, Venezuela and St.

Vincent. A. nephile Taczanewski is known from southeastern United

States, Peru, Brazil, +Guiana, Haiti and Bermuda. We found it on both

Mona and Desecheo. A. trituberculatus Becker has been recorded from

+ Mississippi and Haiti.

Rhomphea is found throughout most of the world’s tropics and sub-

tropics. R. paradora (Taczanowski) was described from Guiana and

has been reported from St. Vincent, but not elsewhere in the Antilles.

Keyserling states that it is probably only a variety of fictiliwm (Hentz)

which is the only known species in the United States and extends from

‘New England to Florida.

Spintharus contains but two species, one confined to Brazil and one,

fiavidus Hentz, which was described from Alabama and is now known

from all the States to Peru and from St. Vincent. It is a yellow, red

and black creature usually found on the under side of the leaves of small

bushes and should be looked for in the Greater Antilles.

Theridion is a large genus which is found throughout the world.

Curiously enough, with the exception of certain cosmopolitan species,

records from the Greater Antilles are almost lacking. We took antil-

fanum Simon, hitherto only known from St. Vincent, by beating branches

in a sea-grape thicket at Dorodo, near San Juan, Porto Rico, and also

in a narrow ravine in the mountains north of Vinales, Cuba. TZ. fron-

deum Hentz is found from Labrador (specimen in our collection) to

Lower California and on St. Vincent. It will probably be found in the

Greater Antilles. We took fuesslyi Simon, formerly known only from

St. Vincent, by sweeping the tall grass and shrubs on the southwest coast,

of Desecheo. Among the wide ranging tropical and subtropical species

are rufipes Lucas, studiosum Hentz, tepidariorum C. Koch, and vituper-

abile Petrunkevitch. We have the first two from the Bahamas but the

only record for any of them from the Greater Antilles is that by Banks

of studiosum Hentz from Haiti. It was described from South Carolina

and Alabama and is known not only from “tropical and subtropical North

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 85

and South America” but also from Bermuda. Finally, volatile Weyser-

ling is recorded from Florida, +Venezuela and (immature specimen)

Culebra Island, Porto Rico.®

Theridula is found in the Mediterranean region, Africa, Madagascar,

tropical and eastern Asia, Japan, Philippines and in America from Lab-

rador (epulenta in our collection) to Peru and Brazil. Simon gives the

distribution of opulenta Hentz as the Mediterranean region, western

Africa and the Transvaal, Madagascar, Ceylon, the Antilles and North

America. It was described from Georgia. There are a number of

synonyms which extend its distribution to Peru and complicate matters.

Banks reported it, under the name of triangularis eyserling from Caya-

mas, Cuba, and from Haiti. He also reported spherula (Hentz), prob-

ably another synonym, from Haiti. He identified the specimens which

we took in Cuba at Cabanas, Cerro de Cabras (near Pinar del Rio),

Banos San Vincente, and Guane as triangularis, and others which we took

at Cerro de Cabras as quadripunctata Keyserling. This latter is prob-

ably also a synonym. Apparently there are no other records for the

Greater Antilles.

Latrodectus is found throughout most of the tropical and subtropical

world, including Madagascar and Australia. The species tend to have

bright red markings and “have been notorious in all ages and in all re-

gions of the world where they occur on account of the reputed deadly

nature of their bite’ (Cambridge). JL. mactans (Fabricius) extends out-

side even the subtropics, being found from New York to Tierra del Fuego.

Cambridge (Proc. Zodl. Soc. London, April, 1902, p. 253) points out that

it is hardly separable from tredecim-guttatus, which is found in the

Canaries, Madeira and the Mediterranean region. The only West Indian

record is by Banks at Mayaguez, Porto Rico and Cockerell in Jamaica.

We took it at Cabanas and Banos San Vincente in Cuba. L. geometricus

C. Koch is found in Cape Verde Islands, Africa, Madagascar, India, Aus-

tralia, Bermuda and tropical South America, the type locality being Co-

lombia. The only West Indian records I have seen are Santiago de las

Vegas and Havana, Cuba.

Petrunkevitch gives the distribution of Teutana grossa (C. Koch) as

“Mexico, Guatemala, Costa Rica, West Indies, Brazil, Uruguay, Chile,

Argentina, Is. Juan Fernandez, (Europe, Africa).” I can find no au-

thority for the West Indies and believe that he mistranslated Simon’s

6 Since writing the above I have heard from Mr. Banks that he has seen Theridion

interruptum Banks and 7’. rufipes Lucas from Havana, Cuba. The former is known

elsewhere only in Florida, but the latter is a cosmotropical species. They are not in

cluded in the discussion at the end of this paper.

86 ANNALS NEW YORK ACADEMY OF SCIENCES

“les iles de l’Atlantique,” although there is no reason for not expecting

it in the West Indies.

Lithyphantes is a cosmopolitan genus which has been recorded on the

American mainland from Canada to Patagonia, but not from the An-

tilles. We took septemmaculatus Kevserling in Cuba in a mangrove

swamp hear Cabanas and im a flower garden at Banos San Vincente in

the mountains. The localities given in the original description are stated

as follows: “Herr Marx fing dieses Thier im Juli bei Denver in Colum-

bia und im December bei Enterprise in Florida.” Doubtless Denver,

Colorado, was intended. It is also recorded from Curacao.

Mysmena is known from France, northern Africa, Cevlon, Philippmes

and United States (two species, both in Florida, one of them being

known also from the District of Columbia). We got several young speci-

mens of the genus from the fallen leaves at the base of the cliffs at Banos

San Vincente, near Vinales, Cuba.

Theridionexus cavernicolus Petrunkevitch is the only species in its

genus and is known only from the Peru Cave in Jamaica. Although he

places this creature among the Theridiide, Petrunkevitch says: “Its gen-

eral appearance, the long front legs and the globose abdomen, and most

of all the presence of a well developed tarsal comb, speak for its close re-

lation to the family Theridiide. On the other hand, the structure of the

mandibles, the shape of the cephalothorax, and especially the presence of

a tibial apophysis in the male palpus, are characters which are found only

in the Argiopide. It is, therefore, impossible to place the genus Theri-

dionexus with sufficient reason in either of these families; it forms a new,

intermediate group.” From the standpoint of phylogeny, perhaps one

might say it belongs to an old intermediate group, and if this be true its

discovery,in a cave in the mountains of Jamaica has an added interest.

LINYPHIID-®

The distribution in America of this family is interesting. For the

most part it is northern, but it has a number of representatives in south-

ern South America and very few between. In addition to certain genera

found only in Greenland or the extreme northern pait of the hemisphere

and others found only in southern Patagonia or Tierra del Fuego, there

are others such as Gonatium and Gongylidiellum which are found at both

extremes but not between. This discontinuous distribution may be due

to faulty taxonomy or there may be natural causes for it. It is, however,

not unknown in other organisms or even in other groups of spiders. This

family is sometimes considered to be a subfamily of Argiopide.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 87

Bathyphantes is known from Europe, Asia, Australia and New Zea-

land, as well as from America. Of the forty American species, thirty-one

are not recorded south of the United States, four are more southern but

not known south of Guatemala, and four are from the extreme southern

part of South America. The final one is the recently described sems-

cincta Banks, collected by us in a flower garden at Bafios San Vincente,

near Vinales, Cuba.

Ceratinella is a palzearctic genus which has one species in this hemi-

sphere. This species, brunnea Emerton, is found from Labrador (speci-

men in our collection) to New York. We also have a specimen of the

genus, species undetermined, from the pine-palmetto plains south of

Pinar del Rio, Cuba.

Ceratinopsis is an American genus with twenty-three species of which

eleven are not recorded south of the United States (except, now, see

anglicana) ; three from Mexico or Central America; and of the nine

South American ones about half are confined to the southern part of that

continent, three of them being known only from the region of Tierra del

Fuego. We took anglicana (Hentz) by beating oak branches in a dense

thicket at about 125 meters elevation on Cerro de Cabras, near Pinar del

Rio, Cuba. We also took the genus, species undetermined, in a ravine at

an elevation of about 300 meters near Banos San Vincente, Cuba. There

are no other records for the genus in the West Indies.

Linyphia is another genus which extends from the northern to the

southern extremes of this hemisphere, in fact it is nearly world wide in

its distribution, but it is rather better developed in the tropics than some

of its relatives. The only West Indian species is coccinea Hentz, which

is found in {Florida and Haiti.

Microneta has a wide distribution, especially in temperate regions.

In America there are twenty-one species confined to northern United

States and Canada; one to Mexico; one (varia Simon) to St. Vincent;

and one to Brazil. If the last two are correctly placed, the genus is

likely to be found in the Greater Antilles.

ARGIOPID

This large family which includes the true orb-weavers is unsatisfactory

material for a study of distribution because of the uncertain limits of

some of the genera. Petrunkevitch and others have dodged the issue by

putting eighteen of them in the Cohors Araneus, and I can only do like-

wise, putting, however, the probable generic name in parentheses. One

of the subfamilies (Linyphiine) into which Simon divides the Argio-

pide has already been considered, treating it as a family.

ANNALS NEW YORK ACADEMY OF SCIENCES

oa)

TETRAGNATHINZ

Cyatholipus is a genus erected by Simon to contain two of his species

from southern Africa and one, dentipes Simon, from Jamaica. It is so

distinct that Simon considers it to be the representative of a special group

of the subfamily. |

Tetragnatha is an almost cosmopolitan genus. The known distribu-

tion of antillana Simon is Mexico, Central America, Porto Rico (Lares),

and St. Vincent. 7. elongata Waleckenzer may be fcund in the Greater

Antilles as, although it is a species of Canada and United States, Walcke-

ner has recorded it from Guadeloupe. It occurs near streams and

usually hangs its web partly, at least, over the water. T’. laboriosa Hentz

has been recorded from Alaska, much of the United States and Porto

Rico (Utuado). Mr. Banks writes me that he has seen a specimen from

Havana. TJ’. piscatortia Simon has been reported only from St. Vincent

but we found it in Porto Rico at an altitude of about 500 meters on

El Duque and also near Arecibo. TY. vicina Simon is recorded from +St.

Vincent and Porto Rico (San Juan). Banks records the genus from

Haiti on the basis of immature specimens and we have a number of such

specimens from the edge of a mangrove swamp near Cabanas, Cuba.

See also the discussion of Eugnatha.

Eugnatha is probably not more than a subgenus of Tetragnatha but,

following Banks and Petrunkevitch, it will be separately considered.

There is one species apparently confined to Mexico and one to northern

United States. HH. pallescens (F. Cambridge) is recorded from +New

York, New Jersey, Florida, Texas, New Mexico, Mexico and Cuba (San-

tiago de las Vegas and Havana). JF. gracilis Cambridge is recorded _

from Mexico, tGuatemala and Bayamon, Porto Rico. We found it at

Naguaba, San Juan and Mayaguez in Porto Rico and at Banos San Vin-

cente in Cuba.

Meta, as construed by Simon and others, is a fairly large and almost

cosmopolitan genus. As is pointed out below, Mr. Banks considers

bigibbosa (Keyserling) to be a Leucauge and he expressed his further

opinion that Meta does not occur in the tropics of America at least.

Alcimosphenus has two species: bifurcatus Petrunkevitch from

Jamaica; and licinus Simon. The latter is recorded from St. Vincent;

Adjuntas, Porto Rico; Haiti and +Santo Domingo; Santiago de las

Vegas, Cuba; and +Jamaica. We found it in Cuba at Cerro de Cabras

and Banos San Vincente.

Leucauge, more widely known as Argyroépetra, is a large genus which

is generally distributed in the warmer parts of the world. JL. argyra

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 89

(Walckener) ranges from California and Florida to Brazil and is recorded

also from Cuba (Havana), Haiti, Porto Rico (Aguadilla, Arroyo, Lares

and El Yunque), Vieques, {Guadeloupe and St. Vincent. We took it at

nearly all stations from Guane to Guantanamo in Cuba, at Hope Gardens

and Montego Bay in Jamaica, on Mona, and at Arecibo and San Juan

in Porto Rico. L. regnyi (Simon) has been recorded only from St. Vin-

cent, although it is possible, as suggested to me by Mr. Banks, that Argy-

roépetra bigibbosa Keyserling, which Petrunkevitch questionably put in

Meta, is a synonym. If so, its range includes also Colombia, Porto Rico

(Aguadilla and El Yunque) and Haiti. In that case bigibbosa is the

prior name. We took it in Porto Rico in the foothills of El Duque near

Naguabo and at Arecibo, on Mona and Desecheo, and very commonly

throughout Cuba. JL. venusta (Walckener) is more generally known

as Argyroépeira hortorwum (Hentz) Emerton. It is fairly common

throughout the United States (Georgia being the type locality) and south

to Colombia. Its recorded insular distribution is Bermuda, Bahamas,

Cuba (Santiago de las Vegas) and St. Vincent. We did not take it in

Cuba but did on Mona and in the mountains south of Arecibo, Porto

Rico. As it is a very striking ‘“‘green and silver-white spider, tinged with

golden, and sometimes with orange-yellow or copper-red spots” and

usually rests in a conspicuous web, it is not likely to be overlooked.

Dolichognatha is found in Ceylon, western Africa, and (the species

given here) tropical and subtropical America. It is closely related to

Diphya which has a somewhat similar distribution: South Africa, Mada-

gascar, and Chile. D. tuberculata (WKeyserling) has been found in ¢Flor-

ida, Mexico, Costa Rica, Venezuela and St. Vincent. It will probably

be found in the Greater Antilles.

NEPHILINA

Nephila is found throughout much of the warmer parts of the world

but seems to be rare in the West Indies, clavipes (Linnzus) being the

only species recorded, although it has gone under a number of names, of

which wtldert, wistariana and concolor, all by McCook, should be men-

tioned. Its known range is from southern United States to Peru and

Brazil, Bermuda, Bahamas, +Jamaica, Santo Domingo and Porto Rico

(Aguadillo). We took an immature Nephila at Cabafias, Cuba, and as

Mr. Banks has seen a specimen of clavipes from Havana our specimen

doubtless belongs to this species. We have a number of adults from

Mona.

90 ANNALS NEW YORK ACADEMY OF SCIENCES

ARGIOPINA

Argiope is found.in the warmer parts of the world, less frequently

in the temperate. Three (aurantia and the two mentioned here) of the

eight American species are wide ranging but the others are more re-

stricted. A. argentata (Fabricius) is said to be found from southern

United States to Patagonia, including all the islands. Among the latter

are St. Vincent, Martinique, Culebra, Porto Rico (San Juan, Utuado

and Aguadilla), Santo Domingo and Isle of Pines. We also have it

from several stations in Jamaica. A. trifasciata (Forskal) is almost cos-

mopolitan. Combining the records of Banks (fastuosa Olivier) and our

captures, it is safe to say that it is found throughout Cuba and Banks

had it from San Juan, Porto Rico, but I know of no other definite rec-

ords from the Antilles. We have a number of undetermined specimens

of the genus from Mona, Desecheo and Porto Rico (San Juan and

Arecibo) which may be one, or both, of these species.

Gea is known from West Africa, tropical Asia, East Indies, Polynesia,

Australia and, by two species, America. One of these is reported only

from Mexico. The other, heptagon (Hentz) has been found in tsouth-

eastern United States as far north as the District of Columbia, in Guate-

mala, Brazil and Cuba (Havana). We took it in Cuba at Pinar del Rio

and Banos San Vincente.

Cyrtophora is found throughout the tropical and subtropical world.

It has three species in northern South America; one in California; and

one, sellata Simon, in Santo Domingo.

Cyclosa has a world-wide distribution. C. caroli (Hentz) is found

from southern United States ({Alabama) to Venezuela; also in St. Vin-

cent and at Havana, Cuba. C. oculata (Walckenzr) was described from

BHurope (France and Italy) but Simon says it is probably of American

origin and introduced to Europe where it is rare and localized. The

American localities given by him are Venezuela and “Antilles.” It would

be interesting to know which of the Antilles are concerned and how

abundant and widespread the species is in this hemisphere. ‘The re-

corded distribution of C. walckeneri (Cambridge) is California, Mexico

to +Colombia, Brazil, Cuba (Santiago de las Vegas) and Haiti. We

took it in Cuba also at Cerro de Cabras near Pinar del Rio.

Edricus is an American genus with a range from Mexico to Peru and

Brazil. We took crassicauda (Keyserling) in the mountains about mid-

way between Arecibo and Utuado, Porto Rico. ‘This species is found on

the mainland from Mexico to +Colombia.

Mangora has a wide distribution but has not been reported from

Africa, Madagascar, the Pacific Islands or Australia. Neither has it

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 91.

previously been reported from the West Indies although it is known to

extend from northern United States to Brazil. We took a common

United States species, placida (Hentz), in the Cerro de Cabras near

Pinar del Rio, Cuba. This is not a species which would be likely to be

carried by commerce, and although the United States Army operated to

some extent in these hills it is not probable that this is a human intro-

duction. If it were we would have expected to find the species nearer

the regular lines of traffic. Alabama is the type locality.

Acacesia foliata (Hentz) is the only species in the genus as construed

by Petrunkevitch. It is recorded from New England to Panama and

from Haiti. It was described from Alabama.

Eustala is a fairly large American genus, chiefly found in Central

America but ranging throughout most of the mainland south of Canada.

FE. anastera (Walckener) has many synonyms, the most common of which

is Hpeira prompta. It was described from Georgia and its distribution

includes most of the United States and southward to Costa Rica and the

Galapagos; Cuba (Cayamas and Havana according to Banks and Guane,

Pinar del Rio, Banos San Vincente and Santiago in our collection), Haiti,

Porto Rico (2800 feet altitude on El Yunque according to Banks and

Arecibo in our collection) and St. Vincent. The variety, or separate

species, conchlea (McCook) is known from +California, +Florida and

Utuado, Porto Rico. The recorded distribution of F. fuscovittata (Key-

serling) is Mexico to Paraguay, Guatemala being the type locality, Cuba,

Porto Rico and St. Vincent. We found it in nearly all parts of Cuba

and Porto Rico which we visited but not on Mona or Desecheo.

Drexelia is probably a synonym of Larinia, a genus found throughout

most of the subtropical, and, more rarely, tropical world. As limited

here (following McCook, Cambridge and Petrunkevitch) there are two

species in America: one in Mexico, and one, directa (Hentz), recorded

from +southeastern United States to Panama and at Havana, Cuba. We

took it in several places on the plain south of Pinar del Rio, Cuba.

Cohors Araneus. As was stated above, this group includes a number

of genera whose limits have not been clearly defined.

A. (Neoscona) arabesca (Walckener) is found on the mainland from

. Labrador to Mexico and also on Curacao. We took it in western Cuba

at Pinar del Rio and Guane by sweeping in meadow land. It was de-

scribed from the Carolinas.

A. (Verrucosa) arenatus (Walckener) was described from Georgia

and is known from New Jersey west to California and south to Panama.

McCook says: “I have specimens collected by the late Mr. W. H. Gabh

from San Domingo varying in but slight particulars from those above

described.”

92 ANNALS NEW YORK ACADEMY OF SCIENCES

A. (Eriophora) balaustinus McCook is recorded from +Florida to

Lower California and Mexico; San Juan, Porto Rico; Haiti and Santo

Domingo; Cayamas and Havana, Cuba; Jamaica; and Swan Island.

The latter island, lying between Cuba or Jamaica and Central America,

is a very interesting locahty. It is to be regretted that we know so little

of its fauna. We took this species on Mona and in a rotten banana trunk

near Arecibo, Porto Rico.

A. (Neoscona) benjaminus (Walckener). lJLabrador (in our collec-

tion) to Guiana and in Martinique. It is probably in the Greater

Antilles.

A. (Epeira) bispinosus (Keyserling) is recorded from +Calitornia,

Arizona, Panama and Haiti.

A, (Singa) crewii (Banks). +aiti.

A, (Singa) cubana (Banks). The type locality is Havana, Cuba.

We took it on the same island at Pinar del Rio and Banos San Vincente

by sweeping the grass at the edge of water.

A. (Neoscona?) granadensis (Keyserling) is recorded from +Colombia

and the Luquillo, Porto Rico. In giving the latter record, Banks remarks

that it is close to trivittata Kevserling which is here considered to be a

synonym of arabesca. !

A. (Epeira) gundlachi (Banks) was described from a specimen taken

by us on the sandy plain about 12 kilometers south of Pinar del Rio,

Cuba.

A. (Epeira) incertus (Cambridge), described from Costa Rica, has

hitherto been known only from Mexico and Central America. We got it

in Cuba at Cabanas and Esperanzia on the edge of mangrove swamps, in

the mountains north of Vinales on oaks, and south of Pinar del Rio on

the palmettoes of the dry sandy plains. A species so catholic in its eco-

logical tastes is doubtless more widely distributed than these records show.

A. (Metepeira) labyrintheus (Hentz) is said to occur “from Labra-

dor to Patagonia, including all the islands.” The only definite Antillean

records I have seen are Barbados; St. Vincent; Culebra and Vieques;

Aguadilla, Porto Rico; and Havana, Cuba. We took it at Mayaguez,

Porto Rico, and on Desecheo. Its type locality is North Carolina.

A. (Verrucosa) mexicanus (Lucas) of which wndecimvariolata Cam-

bridge is considered to be a synonym is known from Panama, Costa Rica,

+Guatemala, Mexico and Santo Domingo.

A. (Neoscona) nauticus (1. Koch) was described from Africa. It

is said to be found throughout the world’s tropics and also in New Hamp-

shire and Tennessee. Its only Antillean records are St. Vincent and

Haiti.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 93

A. (Neoscona) neotheis Petrunkevitch is a name proposed for theisii

of Keyserling and McCook but not theis of Walckener which is a Poly-

nesian species. If this synonymy be followed, the known distribution is

California, Mexico, Guatemala and probably the followimg Antillean lo-

ealities,—Culebra; El Yunque, Bayamon, Aguadilla and Mayaguez,

Porto Rico; and Haiti. In givimg the Porto Rican records Banks says

that “[thetsw| is smaller and more slender than H. oaxacensis Keyser-

ling, and I think different, although small specimens of the latter look

much like large specimens of the former.” See the next species.

A. (Neoscona) oaxacensis ( eyserling) is, as is indicated above, con-

fused with neotheis. Petrunkevitch gives its distribution as Pacific Coast

of United States, +Mexico, Panama and St. Vincent. Banks recorded it

from Santiago de las Vegas, Herradura and Havana in Cuba but made

theist Keyserling a synonym. Specimens taken by us at Cabanas and

Pinar del Rio in Cuba, on Mona, and at Mayaguez, Arecibo, Manati, San

Juan and Naguabo in Porto Rico were identified by Mr. Banks as this

species.

A. (Epeira) pegnia (Walckener) is recorded from Colombia, Costa

Rica and most of the United States. Mr. Banks has written me that he

has seen Hpeira globosa Weyserling, here considered to be a synonym,

from Havana, Cuba.

A. perplexus (Walckener) was described as an FP peira in 1842 from

Brazil and Santo Domingo but has, apparently, not been recognized since.

A. (Epeira) pratensis (Hentz) is recorded from most of the United

States but not elsewhere. We took it near Banos San Vincente, Cuba,

and Mr. Banks informs me that he has seen it from Havana.

A. (Epeira) sericatus Clerck apparently has a number of synonyms.

Among them is vulgaris Hentz, the name which Mr. Banks gave to our

specimens from Cabanas, Pinar del Rio and Guane, Cuba, as well as to

those he had from Santiago de las Vegas and Havana in the same island.

In all cases they were about houses and may have been introduced. The

only other American records are from the continent north of Mexico. It

is also found in + Europe.

A. (Marxia) stellatus Walckener was described from southeastern

United States and is found from Labrador (specimen in our collection)

to Guatemala. We took it on the plain ten kilometers south of Pinar del

Rio, Cuba, at the edge of a swampy area.

A. (Wagneriana) tauricornis (Cambridge) was described from a

number of localities in Guatemala and Panama. It is recorded also from

Colombia, Mexico, Louisiana, Alabama, Florida, Cuba (Santiago de las

Vegas) and Haiti.

94. ANNALS NEW YORK ACADEMY OF SCIENCES

A. (Wagneriana) undecimtuberculatus (Keyserling) is known from

tColombia, Panama, Guatemala, Mexico, Florida, Cuba (Santiago de las

Vegas) and Haiti.

A. (Eriophora) variolatus (Cambridge) is found in southern and

western United States, Mexico, +Guatemala, Venezuela and St. Vincent.

It is probably in the Greater Antilles also.

A. (Metazygia) wittfelde (McCook) is known from +Florida, Mexico,

the Bahamas and Havana, Cuba. We took it at Mayaguez, Porto Rico,

in a hotel. ;

Gasteracantha is widely distributed in the tropical and subtropical

regions of the world. G. cancriformis (Linneus) is found from North

Carolina and California to Paraguay; the Bahamas; El] Guama, Santiago

de las Vegas and Havana, Cuba (according to Banks under the name of

hexacantha Fabricius) ; Haiti and Jamaica. We have it from the Ba-

hamas, Jamaica, Mona and Banos San Vincente in Cuba. G. hilaris

Thorell is recorded from ¢St. Bartholomy, Porto Rico (Aguadillo and

Adjuntas), and Haiti. In Porto Rico we found the under side of leaves

of young coco palms at San Juan and also in the mountains south of

Arecibo festooned with its webs. Banks has referred G. canestrinti Cam-

bridge to this species, thus extending its range to Antigua and Dominica,

but Petrunkevitch does not follow him in this. G. sexserrata (Walek-

ener) is recorded from +Cayenne, Haiti and Porto Rico (Bayamon). I

took it several years ago on Key Largo, Florida, so it is probably in Cuba

also. G. tetracantha (Linneus) is recorded from California (if pallida

Koch be a synonym), Haiti, Culebra, St. Thomas, St. Vincent, Guade-

loupe and three of the Grenadines. We can now add to this list. Porto

Rico (near Arecibo) and Desecheo.

Micrathena is the Acrosoma of authors. As now constituted it con-

tains more than 125 species, all American, and the great majority trop-

ical. M. armata (Olivier) is known only from + Hispaniola and Jamaica.

M. cubana (Banks) is recorded only from +San Diego de los Baftos and

possibly Havana, Cuba. We took it in Cuba at Banos San Vincente.

M. flavomaculata (Keyserling) is known only from Haiti; and forcipata

(Thorell) only from Cuba. M. horrida (Taczanowski) is recorded from

Mexico, Peru, +Guiana, Brazil and Jamaica; obtuspina (Keyserling)

from + Mexico to Brazil and also from Porto Rico. Finally, rufopunctata

(Butler) and sloanei (Walckenzr) are known only from Jamaica. It is

decidedly curious that there are no records of the genus in the Lesser

Antilles.

Glyptocranium is strictly an American genus. The related Old World

genera are found in Australia, Africa and southern Asia. It has two

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 95

species which are found only north of Mexico (one ranging even to

Alaska), two in Mexico and one, gastercanthoides (Nicolet), in Brazil,

tChile and Jamaica.

Epecthinula minutissima Simon is the only species of its genus and

is known only from Jamaica. The genus is related to Hpecthina, which

lias but a single species, found only in Venezuela. According to Simon,

there are three other genera belonging to the same group of the Argiop-

ine. One of these is found only in northern South America; one in

South Africa and western Australia; and one in Algeria, New Caledonia

and northern South America. |

MIMETID

This is a small, for the most part tropical, family the members of

which are usually found near or on the ground and make no definite web.

None have heretofore been recorded from the West Indies.

Mimetus is found in the Mediterranean region, Africa, India, New

Zealand and America. In the latter hemisphere there are nine species,

of which two are recorded only from South America, five from Mexico

and Central America, one from North Carolina. The remaining one,

mterfector Hentz, ranges over much of United States and we took it on

the sandy plain south of Pinar del Rio, Cuba. The genus occurs also in

Porto Rico, for we took’an immature specimen of it near Arecibo.

THOMISID®

These are called crab-spiders, not only on account of their shape but

also because they usually walk sidewise or backward. They spin no

snares but catch their prey either by pursuit or by Jying in wait for it,

being much favored with concealing colors. It is a large family the

American members being chiefly found in or near the tropics although

it ranges from Greenland to Patagonia.

MISUMENINZ

Misumenops is a split, possibly unwarranted, from the cosmopolitan

Misumena, to include a number of American species. Jf. americanus

(Keyserling) is recorded from United States, Guatemala and St. Vin-

cent. M. asperatus (Hentz) ranges from Canada to Costa Rica and is

known also from Cuba, Haiti, Porto Rico (Bayamon and El Yunque)

and St. Vincent. We found it very common throughout Cuba and took

it also on Mona and at Mayaguez and San Juan, Porto Rico. J. bellulus

(Banks) is recorded only from Florida but Mr. Banks has written me

96 ANNALS NEW YORK ACADEMY OF SCIENCES

that he has it from Havana, Cuba. J/. celer (Hentz) probably includes

M. spinosa Keyserling. Mr. Banks has recorded spinosa from Santiago

de las Vegas and Cayamas, Cuba, and has named the specimens which

we took at nearly all of our stopping places in that island from-Guane

to Guantanamo spinosa, while he named the specimens which we took at

Mayaguez, Arecibo, Manati and Naguabo in Porto Rico celer. On the

mainland, celer is found from Massachusetts to Mexico. ‘The known dis-

tribution of oblongus (Keyserling) is from Massachusetts to Illinois and

south to Georgia and New Mexico; also in Cuba (Santiago de las Vegas).

M. viridans (Banks) is recorded only from Florida but Mr. Banks has it

from Havana, Cuba. See also the next genus.

Misumessus echinatus Banks was described from material beaten

from oak trees on Cerro de Cabras near Pinar del Rio, Cuba. According

to the system followed here, it should probably be put under Misumenops.

STEPHANOPSINA

Isaloides contains but two species: one from Mexico and one, tous-

saintu Banks, from +Haiti and Cuba. It is closely related to and con-

sidered by Simon to be not more than a sub-genus of Hedana. He would

also include the South American genera Diea and Erissus. This adds

interest to the distribution data, one section of the genus being found

only in Hispaniola and South America while the other section is known

only from Ceylon, southwestern Asia, Philippines, Malasia, the Tonga

Islands, Australia and New Zealand.

Stephanopsis is confined to Madagascar, Malasia, Melanesia, Poly-

nesia, Australia and America. In America there is one species in Pan-

ama, eight in South America (chiefly Chile) and one in Tortola of the

Virgin Islands. Banks has recorded the genus from Santiago de las

Vegas, Cuba. He states that he had two species, both represented only

by Immature specimens, and suggests that one of them is a young

pentagona \Keyserling. This species is now considered to belong to

Onocolus. ‘The species is known from Panama, Peru and Brazil. and as

Onoculus is solely a South American genus the interest in the distribu-

tion of Stephanopsis is not lessened.

Tobias is an American genus, four species being confined to northern

South America and one, rugosus (Taczanowski), being recorded from

Brazil, +Guiana, Peru, Central America and Haiti.

PHILODROMINZA

Tibellus is widely distributed in the temperate regions of the world,

less so in the warmer regions. In America there is one species confined

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 97

‘

to each of Mexico, Guiana and Paraguay while the fourth species, ob-

longus (Walckener), occurs in +Hurope, Asia, Alaska and most of United

States. We took the latter species on the dry plain nine kilometers south

of Pinar del Rio, Cuba.

CLUBIONID-®

The Clubionids are frequently confused with the Drassids and their

habits are much the same. They make nests in rolled leaves, under

stones, or in rubbish. ‘The alphabetical arrangement of genera used by

Petrunkevitch is especially confusing in this family as there are very

distinct divisions which may be of family rank. The arrangement of

Simon will be followed here.

SELENOPINZ

Selenops is the only genus. It is found throughout most of the

world’s tropics. They are flat creatures which crawl under bark or stones

whence they dart out for their prey. S. atssus Walckener is known from

Bahamas, Tortugas, Cuba (Cayamas and Santiago de las Vegas) and

+Martinique. We found it hiding back of the boards of a house at

Cabanas, Cuba. Macleay in describing S. celer (Macleay) said that

it is common in Cuba. We took it north of Vinales, Cuba, on banana

trees. It is recorded only from this island and Buen Ayre, off the coast

of Venezuela. S. insularis Keyserling is known from +Porto Rico (San

Juan, at least) and Haiti. We took it on Desecheo in a rotten log and

also under fallen leaves in a sea-grape thicket.

SPARASSINA

Heteropoda is also found throughout most of the world’s tropics.

There are only four American species: three confined to South America

and the cosmo-tropical venatoria (Linneus). The known West Indian

distribution is Jamaica, Cuba (apparently throughout), Haiti, Porto

Rico (San Juan) and St. Lucia. It is frequently brought north in fruit.

Olios is likewise a cosmo-tropical genus. O. antiguensis (Keyserling)

is known from Haiti, Porto Rico (Utuado), Culebra and +Antigua. 0.

bicolor Banks was described from specimens which we took on Desecheo,

Mona, and at San Turce near San Juan, Porto Rico. O. maculatus

(Blackwell) is reported from Brazil and the “Antilles.”

Pseudosparianthis is an American genus with one species in Mexico,

two in Brazil, one in St. Vincent, and one, cubana Banks, in Cuba

(+Havana and +Santiago de las Vegas).

98 ANNALS NEW YORK ACADEMY OF SCIENCES

CLUBIONINZA

Anyphena in America is largely northern but it does extend even to

Patagonia. It is found also in Japan, the mountains of India and the

western Mediterranean region. We took immature specimens of it near

Cabanas, Pinar del Rio and Guantanamo in Cuba. A. perpusilla Banks

is known only from Santiago de las Vegas, Cuba. A. striata Becker was

described from Mississippi and is elsewhere known only by Banks’s record,

under the genus Aysha, from Santiago de las Vegas, Cuba. He also

records under the same generic name A. velox Becker from Havana, Cuba.

It is known also from + Mississippi, Florida and the Bahamas. A. graci-

lipes Banks is known only from Haiti.

Aysha is a small tropical American genus closely related to the pre-

ceding one. Nine species are found in the region from Central America

to Brazil; one is confined to the Galapagos Islands; two, ferox Simon and

ravida Simon, are recorded only from Santo Domingo; and, finally,

tenuis (lL. Koch) is known from Cuba, + Haiti, Santo Domingo, Porto

Rico (San Juan), Culebra and St. Vincent. We took tenwis on Desecheo

and obtained immature specimens of the genus on Mona.

Chiracanthium inclusum (Hentz) was reported by Banks from San-

tiago de las Vegas, Cuba and Aguadilla, Porto Rico. We took it at

Guane and at Guantanamo, Cuba, but not between these two places. We

also took it in Porto Rico near Quebradillas, in the seed pods of Crotolaria

retusca which had been eaten out by Lepidopterous (Utetheisa) larve,

and near Dorodo. It has been found in St. Vincent, Haiti, throughout’

much of tUnited States and in Mexico. The genus is nearly world-wide

im its range.

Clubiona-is another cosmopolitan genus but it is more abundant in

temperate than in tropical regions. In America it extends from Lab-

rador to Patagonia. Its best South American development is in Chile.

(’. maritima I. Koch is reported from Santiago de las Vegas, Cuba, and

+St. Thomas. C. pallens Hentz occurs on the Atlantic Coast from Massa-

chusetts to tAlabama, and we found it in a weedy meadow on Cerro de

Cabras, Pinar del Rio, Cuba.

Eutichurus is reported only from Central and South America except

for insulanus Banks, which is in Cuba, Haiti and +Bermuda.

Oxysoma has ten species in South America. As they are chiefly in

the southern and western part, it is interesting that the eleventh (cubana

Banks, omitted in Petrunkevitch’s catalogue) should be found in Cuba.

It was described from one male from Havana and has not been recorded

elsewhere.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 99

Wulfila is an American offshoot of Anyphena. It is known only from

Mexico, Central America, Bahamas and West Indies. There is one

species recorded only from St. Vincent. W. immaculata Banks was de-

scribed from specimens collected by us at the edge of a mangrove swamp

near Cabanas, Cuba; on the dry coastal plain of Mona; and at an alti-

tude of about 500 meters on El Duque, near Naguabo, Porto Rico. W.

pretiosa Banks was described from a specimen which we took in the Rio

Seco woods on the San Carlos Estate near Guantanamo, Cuba. W.

parvula Banks is confined to Haiti and tenwissima Simon to Jamaica.

CTENINA

Ctenus is a cosmo-tropical genus. America has a large number of

species but apparently none of them live north of southern United States

and the Bahamas. St. Vincent and St. Lucia each have a peculiar

species; haitiensis Strand is known only from Haiti and malvernensis

Petrunkevitch only from Jamaica. We have the latter from Montego

Bay, Jamaica, and unidentified specimens of the genus from Cuba (Banos

San Vincente, Cerro de Cabras and Guantanamo). Banks recorded an

immature Ctenus which “looks very much like Ct. hibernalis Hentz”

from Santiago de las Vegas, Cuba. This species is known from Alabama,

New Mexico, Mexico and the island of Buen Ayre. He also recorded a

female Clenus (Microctenus) which appears to be new from a cave near

Pueblo Viejo, Porto Rico.

Cupiennius is a South and Central American genus except for one

species which comes as far north as Florida and cube Strand which is

known only from Cuba.

It is interesting that, with the exception of widely distributed Ctenus,

enly one genus of this division (Ctenez) of the Ctenine is found outside

of the middle portion of America. That is Uliodon, which is known only

from Madagascar, New Zealand and Australia. |

LIOCRANING

Syrisca is found in Africa and America. There is one species in each

cf Colorado, Utah and Texas; two in South America, and two in the

Greater Antilles. S. insularis (Lucas) is known only from Cuba and

keyserlingt Simon (= Teminius insularis, Keyserling) only from + Haiti

and Santiago de las Vegas, Cuba.

MICARIINZ

Castaneira occurs in the western Mediterranean region, Africa, central

Asia and America. It appears to be better developed in United States

100 ANNALS NEW YORK ACADEMY OF SCIENCES

and Mexico than it is farther south. The only West Indian record I

have seen is a species confined to St. Vincent. We took a common

United States species, descripta (Hentz). in a valley near Banos San

Vincente, Cuba.

CORINNINZA

Corinna is known from Africa, tropical Asia, Malasia and America.

It is well developed in this hemisphere but, although there are eight spe-

cies in St. Vincent which are not known elsewhere, only two species have

been recorded from the remaining Antilles. C. gracilipes (Keyserling)

is known from Havana, Cuba, and from +Haiti. C. humulis (Keyserling)

has been recorded from Haiti, Porto Rico (Luquillo and Hacienda Es-

paranza) and +St. Kitts; Mr. Banks has written me that he has seen a

specimen from Havana, Cuba; and we have a specimen from Mayaguez,

Porto Rico, which is probably this species.

Trachelas occurs in the Mediterranean region, Africa, Madagascar,

India, Malasia and America. In this hemisphere it appears to be best

developed in Central America, but bicolor Keyserling is the only species

recorded from the Greater Antilles. It is known only from Cayamas and

Santiago de las Vegas, Cuba, and from + Haiti.

AGELENID ©

These are popularly called funnel-web spiders because many of the

species spin flat webs which have a funnel-like passageway to their re-

treat. In America the family is best developed in the north and prob-

ably next best developed in the extreme south, being rather weak in the

tropics except in the more mountainous regions.

Hahnia occurs in Europe and the Mediterranean region, the moun-

tains of India, Sumatra, Japan and America. There are seven species

not found south of the District of Columbia, one of them being known

only from Greenland ; two species are in Patagonia and Tierra del Fuego;

and one, ernstt Simon, recorded only from St. Vincent, but taken by us

in a rotten banana stump at about 500 meters elevation on El Duque,

near Naguabo, Porto Rico.

Tegenaria is found throughout most of the cooler and, more rarely,

in the warmer portions of the world. T. domestica (Clerk), better known

as derhami (Scopoli), is.said to inhabit the dwellings of man in all re-

gions of the world, but I know of no definite record from the Antilles.

T. insularia Walckener is known only from Cuba and has not been re-

corded since its description.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 101

PISAURID

The females of these, like those of the Lycosids, carry their egg sacs

about with them. ‘Some species also build a “nursery” for the newly

hatched young, but construct no snare.

Thanatidius has one species in the Amazon region. The only other

one known is dubius (Hentz), which has been reported from +North

Carolina, Alabama, Florida and Cuba (Havana).

Thaumasia, as now constituted, is found only in the warmer parts of

America. TJ. marginella (C. Koch) Simon is usually placed in Dolo-

medes and is recorded from Colombia, Brazil, Jamaica, Haiti, Vieques

and possibly Porto Rico. We found it at Cabanas, Pinar del Rio and

Banos San Vincente, Cuba. Mr. Banks has written me that he has seen

it from Havana, Cuba. As he considers this species to be a Dolomedes,

it may be that the immature specimens taken by us at Pinar del Rio and

identified by him as Dolomedes belong to this species. As now construed,

there are no records for Dolomedes in the Antilles.

LycosI1pz&

The relatively poor development in the Antilles of this fairly large

family of “ground spiders” seems to accord with the distribution of

Coleoptera (see Leng and Mutchler, 1914), among which the ground

forms are more poorly represented than the arboreal ones. Bates, in his

“Naturalist on the River Amazons,” says: “It is vain to look for the

Geodephaga, or carnivorous beetles, under stones, or anywhere, indeed, in

open, sunny, places. The terrestrial forms of this interesting family,

which abound in England and temperate countries generally, are scarce

in the neighborhood of Para—in fact, I met with only four or five spe-

cies; on the other hand, the purely arboreal kinds were rather numerous.

The contrary of this happens in northern latitudes, where the great ma-

jority of the species and genera are exclusively terrestrial. . . . The

remarkable scarcity of ground beetles is doubtless attributable to the

number of ants and Termites which people every inch of surface in all

shady places and which would most likely destroy the larve of Coleop-

tera. These active creatures have the same functions as Coleoptera, and

thus render their existence unnecessary. The large proportion of climb-

ing forms of carnivorous beetles is an interesting fact, because it affords

another instance of the arboreal character which animal forms tend to

assume in equinoctial America, a circumstance which points to the slow

adaptation of the Fauna to a forest-clad country, throughout an immense

lapse of geological time.” The last suggestion, in a modified form, seems

102 ANNALS NEW YORK ACADEMY OF SCIENCES

to be important. It may be that the fauna of “equinoctial America” is,

to a large extent, a relic of past ages; that it arose in a forest-clad earth

and has been driven before the more successful ground-inhabiting forms,

not that it has been slowly developing from ground-inhabiting forms in

the region in which it is now found.

Lycosa is a large genus of world-wide distribution. Several attempts

have been made to split it up, with varying success. One of these puts

cinerea (Fabricius) in the genus Arctosa. This species has been found

in Europe, throughout United States, in Mexico and at El Guama, Cuba.

L. atlantica Marx was known only from Bermuda, but Mr. Banks writes

that he has seen it from Havana, Cuba. JL. aussereri (Keyserling) is

known from ¢Colombia, Vieques and Culebra; L. badia (Keyserling) from

Central America and Cuba; L. fusca (Keyserling) from Central Amer-

ica, Cuba (Havana, according to a letter from Mr. Banks) and Porto

Rico (San Juan); Ll. punctulata Hentz, from United States east of the

Rockies and, according to a letter from Mr. Banks, Havana, Cuba. L.

insularis Lucas is known only from Cuba. We took it there at Guane,

Pinar del Rio, Banos San Vincente, Cabanas and Guantanamo. Banks,

who now considers it to be a Pardosa, recorded it from Santiago de las

Vegas.

Pardosa is likewise of world-wide distribution. In America it is best

developed in the north. There are a few species in South America, but

unless insularis be included the only West Indian species is portoricensis

Banks, which was described from San Juan, Porto Rico. We took it ina

inarsh at San Turce near its type locality.

These two genera, while much confused in the literature, are placed in

different divisions of the family by Simon. Both are too generally dis-

tributed to be of a great deal of interest in a study of distribution.

OxYOPIDZ

The members of this family run about the vegetation with great agility

in chase of their prey. There are only eight genera, of which six occur in

America. Of the remaining two, one is confined to Madagascar and one

io India and Malasia. Two of the six American genera have not been

reported from the Antilles; one of them has but two species and is con-

fined to Brazil, and one has but one species, being confined to Cayenne,

as far as is known.

Hamataliva has a wide tropical and subtropical range but in America

there are only four species known: two in Brazil, one in Mexico and

grisea Keyserling, which is reported from Lower California, southern:

United States. Cuba (Santiago de las Vegas) and Haiti.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 103

Oxyopeidon is found in tropical east Africa, India, Indo-China and

America. With us it seems to be confined to Mexico and Central America

(sIx species) except for rana Simon, which has been known only from St.

Vincent. We found this species fairly abundant on low vegetation on

Mona.

Oxyopes is a nearly cosmo-tropical genus which extends into more

temperate regions. In America it is best developed in Mexico and Cen-

tral America, where there are numerous species. O. pallidus (C. Koch)

was described from the West Indies, but the only definite locality record

seems to be in Walckener’s description of a male from Cuba. O. salticus

Hentz is known from New York to +Alabama, Kansas, California and

south to Bolivia, also in Bermuda, all the Greater Antilles and St. Vin-

cent. It seems to be common throughout Cuba. We found it on Mona

and near San Juan, Porto Rico. In Porto Rico it is also reported from

El Yunque and Culebra.

Peucetia is also found throughout most of the tropical and subtropical

regions of the world. P. poeyi (Lucas) is known only from Cuba, but

viridans (Hentz) is distributed in America from +North Carolina to Cali-

fornia and south to Costa Rica. It is recorded from Cuba (Santiago de

las Vegas and Cayamas), Haiti and Jamaica. We have it from Kings-

ton, Jamaica; Guane, Banos San Vincente, Oriente and Guantanamo in

Cuba; and Mayaguez, Porto Rico. ‘It has, apparently, not been found in

the Lesser Antilles.

SALTICID-®

The “jumping spiders” make no snare but run about freely on the

ground and on the vegetation in pursuit of their prev. The family is

a large one and contains many species still to be described. while the

taxonomy of the known forms is in a far from satisfactory shape. Our

own West Indian collection was in the hands of Mr. Peckham and only

partly worked up when he died. The following records, therefore, do

not entirely cover the material which we have on hand.

Agobardus anormalis Kevyserling is the only species in its genus.

The type specimens are labeled U. S., but Petrunkevitch, following

Peckham and Banks, states that it is probably from the West Indies.

Bythocrotus cephalotes (Simon) is not only the only species of the

genus but the genus is the only one in Bythocroteew, one of Simon’s

divisions of the Salticide. It is not known outside of Haiti.

Compsodecta is an American genus with one species in Guatemala

and two, namely, albopalpis (Peckham) and grisea (Peckham), in

Jamaica. The genus belongs to Simon’s group Pensacoleew which con-

104 ANNALS NEW YORK ACADEMY OF SCIENCES

tains but two other genera, one of which is best represented in Brazil

and Ecuador but extends to Mexico and the other is confined to tropical

western Africa. ;

Corythalia is a fairly large tropical American genus. ‘Two species,

major (Simon) and sellata (Simon), are mentioned in Simon’s “Histoire

Naturelle des Araignees,” II, pages 655 and 649, in such a way that

Petrunkevitch was led to credit them to the West Indies. and, while this

seems to me questionable, I can find no more definite reference. C.

metallica (Peckham) is reported only from St. Vincent. Although it

does not properly come within the scope of this paper, it is worth men-

tioning that we have it from Dominica. C. elegantissima (Simon) is

known only from Santo Domingo, and locuples (Simon) from both parts

of Hispaniola.

Stoidis is also an American genus. It contains but two species at

present but specimens taken by us in Cuba were marked by Mr. Peckham

as new species. 8S. aurata (Hentz) is known from +South Carolina and

Florida. Mr. Banks has informed me that he has seen it from Havana,

Cuba. S. pygmea (Peckham) is reported only from St. Vincent but

we took it in an epiphyte, Ti//andsia utriculata, on Mona.

These two genera, Corythalia and Stoidis, are the only American rep-

resentatives of Simon’s Zenodorew. The other four genera are from

Australia and the Pacific Islands.

The Cohors Dendryphantes of Petrunkevitch includes, in addition to

some unplaced species which should probably be put in the genus

Dendryphantes. the following genera: Metaphidippus, Paraphidippus,

Parnenus and Phidippus. Although this cohors is in large part the

genus Dendrypiantes Koch as construed by Simon it is not very satis-

factory for a study of distribution. The following list is arranged ac-

cording to Petrunkevitch.

Dendryphantes armatus Banks (omitted by Petrunkeyitch) was de-

scribed from Santiago de las Vegas, Cuba, and has not been reported

since.

D. (Phidippus) audax (Hentz) is known from Canada, throughout

the United States and from Santiago de las Vegas, Cuba. _

D. (Paraphidippus) aurantius (Iucas} is found from southern

United States to Costa Rica and in Santo Domingo.

D. (Metaphidippus) capitatus (Hentz). Canada to Mexico and

Santiago de las Vegas, Cuba (in Banks’s list under name of octavus

Heniz).

Dendryphantes mendicus (C. Koch) is credited to “West Indies”

but no definite data is given.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 105

D. (Phidippus) miniatus (Peckham). The known range is Virginia

to Texas and at Santiago de las Vegas, Cuba.

D. (Phidippus) octopunctatus (Peckham) is reported only from

Missouri but I took it in Dominica, the identification being by Peckham.

It is, therefore, probably in the Greater Antilles.

D. (Metaphidippus) proximus Peckham was described from Cuba.

Banks lists it from Santiago de las Vegas and we took it at Santiago de

Cuba, Cristo, Zaza del Media, Cabanas, Cerro de Cabras and Banos San

Vincente, Cuba. We have it also from Kingston, Jamaica.

D. (Metaphidippus) prudens Peckham is known only from Kingston,

+ Jamaica.

D. (Phidippus) regius (C. Koch) is known only from Cuba. Banks

reports it at Santiago de las Vegas and we took it at Guane, a number

of places in the vicinity of Pinar del Rio, north of Vinales (Banos San

Vincente and Merciditas), Cabanas and Zaza del Media. All these

localities are in central and western Cuba.

D. (Metaphidippus) taylori Peckham is known only from Jamaica.

Evophrys is found in Europe, Africa, Japan and America. There are

about fifty American species, the genus ranging from New Hampshire

to Patagonia. One species is found in St. Vincent, another in St.

Thomas; it may, therefore, be found in the Greater Antilles but there

do not seem to be any records as yet. The only other genus of Evo-

phrydez is confined to Central America and northern South America.

Eustiromastix has seven species; three are confined to Brazil, one to

Colombia, two to St. Vincent and haytiensis Banks to Haiti.

Hasarius adansoni (Adouin) is found throughout much of the world’s

tropics but I know of no West Indian records. It is the only American

representative of the genus with the possible exception of bellicosus Peck-

ham from Guatemala.

Hyctia is recorded only from Europe and United States. One of the

three American species, pike Peckham, ranges from New England to

New Mexico. We took it in Cuba at Cabanas, on the plains south of

Pinar del Rio, and at Guane. Mr. Banks has written me that he has

seen it from Havana.

Icius is given by Simon as being found in Europe, Africa and Asia

but he does not give America. We have thirteen species, all found north

of Mexico except one which is confined to Uruguay, one to Florida and

the Bahamas, and seperatus Banks which is known only from Haiti.

Lyssomanes is a large American genus ranging from southern United

States to Brazil. The. Lyssomanee includes seven genera, of which two

are confined to America (the other one being found from Central America

106 ANNALS NEW YORK ACADEMY OF SCIENCES

to Guiana), two to Madagascar, one to India and Ceylon, one to the

Philippines, and one is found in western Africa, Madagascar, the Sey-

chelles, Cevlon, Burma and Indo-China. The group is evidently an an-

cient one. We have an undetermined specimen of Lyssomanes from Porto

Rico. JL. antillanus Peckham is known from both parts of Hispaniola

and we have it from Jamaica (Lapland). JL. viridis (Walckener), the

most northern species of the genus, is known from tsouthern United

States, Central America, Haiti and (by letter from Mr. Banks) Havana,

Cuba.

Marpissa occurs in Europe, Asia including Japan, and America. On

the occidental mainland it ranges from northern United States to Brazil

but the only Antillean record is incerta Koch from St. Thomas.

Menemerus bivittatus (Dufour) = melanognatha (Lucas) is almost

cosmopolitan if the svnonymy of Peckham and Petrunkevitch be ac-

cepted. Its only Antillean record, however, seems to be Santiago de las

Vegas, Cuba. We have it from about plantation buildings at Cabanas,

Cuba and also from Culebra.

Metacyrba is considered by Simon, but not by Petrunkevitch, to be a

synonym of Fuentes. In any case it is solely American, and it belongs

to a group, Mzvivee, which is known only from America. The only

Antillean species is pictipes Banks from Haiti.

Myrmarachne is practically cosmopolitan. 1. melanocephalus Mac-

Leay is said by Lucas to be from Cuba and Petrunkevitch so records it

without further comment, but the original description says it “is a native

of Bengal and I present a figure of it, made by my friend Mr. C. Curtis,

in order to show the relation which it bears to the American subgenus,

called Myrmecium by Latreille.” M. parallelus (Fabricius) has only the

unsatisfactory record of “+Antilles.”

Nilacantha cockerelli Peckham is the only species of its genus. It is

found in Jamaica and Haiti. The Thiodinz, to which it belongs, is an

American group.

Peckhamia has but four species. It ranges, on the mainland, from

Canada to Panama. A related genus, forming with it the group Peck-

hamiez, is only known from Brazil. There are no records from the

West Indies except for an immature specimen from Haiti, but Mr. Banks

informs me that he has seen a specimen of the United States species,

P. picata (Hentz), from Havana and we took an immature specimen of

the genus at Cerro de Cabras near Pinar del Rio, Cuba.

Pellenes is found in Europe and the Mediterranean region, north-

eastern Asia and America. In the latter region there are more than

fifty species but it is not known south of Central America. There are

—

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 107

but two West Indian records: banks: Peckham and translatus Peckham,

both confined to Jamaica. We took specimens of the genus which Mr.

Peckham marked as new species at Cristo and Pinar del Rio, Cuba, and

we also took coronatus (Hentz) at Pinar del Rio. This species ranges

from New York to Mexico.

Plexippus is found throughout most of the world but especially in

the warmer regions. P. paykulli (Adouin) is a cosmo-tropical species

which has been recorded from Santiago de las Vegas and Havana, Cuba;

Haiti; Lares, Adjuntas, San Juan and Arecibo, Porto Rico, and St.

Vincent. We took it at Guane, Cuba.

Prostheclina Keyserling is considered by Simon to be synonymous

with Saitis. This is probably correct but as there is some doubt they

will be kept separate here except that, in giving the general distribution

of Saitis, Prostheclina will be included. In America Prostheclina seems

to be limited to the Greater Antilles. P. parvula Banks is known only

from Cayamas, Cuba. P. perplexides Strand, venatoria Peckham, and

viarva Peckham are known only from Jamaica (+Ipswich, +Port Antonio

and +Moneague, respectively). P. morgani Peckham was described from

Kingston, Jamaica, and perplera Peckham from Mandeville, Jamaica.

Both species are reported also from Haiti. P. illustris (C. Koch) was

described from Porto Rico and has since been reported by Banks from

an altitude of 2000 feet on E! Yunque in that island. P. signata Banks

is listed in the original description from Utuado and Aguadilla, Porto

Rico; also from Culebra. He has since reported it from Haiti and we

took it at San Turce and Dorodo, both near San Juan, and in the

mountains south of Arecibo, Porto Rico, as well as on Desecheo. |

Saitis, including Prostheclina, is known from southwestern Europe,

western and southern Africa, India, Malasia, Polynesia, Australia and

America. There are three species in South America; the others are from

the Greater Antilles. With the exception of those mentioned under

Prostheclina, all the known Antillean species are confined to Jamaica,

namely, anne Cockerell, +Kingston; defloccatus Peckham, +Kingston,

and inutilis Peckham, no definite locality given.

Synemosyna is an American genus which, with Simonella from the

American tropics, forms the group Synemosynex. Two species of Syne-

mosyna are found in Brazil, one in St. Vincent, and the United States

species, formica Hentz, is listed from Cayamas, Cuba.

Thiodina is an American genus which ranges farther north than the

other members of the American group, Thiodiner. T. sylvana (Hentz)

is known from tSouth Carolina to Panama and, under the name of retia-

_ rlus Hentz, from Cayamas, Cuba.

108 ANNALS NEW YORK ACADEMY OF SCIENCES

Wala is an American genus. W. peckhami (Cockerell) is known

trom +Jamaica and Haiti. We have it from Montego Bay, Jamaica, and

Banos San Vincente, Cuba. JW. vernalis (Peckham) has been reported

from Santiago de las Vegas, Cuba; Jamaica; Haiti; San Juan and Agua-

dilla, Porto Rico; Vieques; +St. Vincent and Bermuda. We took it on

Mona and Desecheo; and at Quebradillas, Manati, Dorodo, San Turce

and Naguabo, Porto Rico. We also took undetermined specimens of the

genus in the vicinity of Pinar del Rio, Vifiales, Cabanas, Cristo and

Guantanamo, all in Cuba.

Zygoballus and a Brazilian genus, together forming the group Zygo-

ballee, are American. It ranges from Panama northward. The only

record for the West Indies is swavis Peckham from Mandeville, Moneague

and Kingston, t+Jamaica. We have it from Montego Bay, Jamaica. We

also have a specimen of the genus, marked by Peckham as a new species,

from Cerro de Cabras, near Pinar del Rio, Cuba.

DISTRIBUTION OF FAMILIES

There are twenty-three families of Araneze now known in the Greater

Antilles. Five of these (Dictynide, Cicobiide, Oonopide, Dysderide and

Mimetide) are added by this list. One family, Leptonetide, has three

species, two genera, on St. Vincent, but it has not yet been found in the

Greater Antilles, although it occurs on the mainland from California to

Brazil. It is placed by Simon between the Sicariide and Oonopide. Its

species are small, long-legged creatures for the most part cavernicolous,

and should be looked for in the Greater Antilles. It is fairly widespread

but is not yet known from southern Africa, Madagascar, Australia or the

southern Pacific Islands.

There are eleven small families which are found on the mainland but

have not, as yet, been reported from any of the Antilles. The Atypide

(p. 77) and Hypochilide (p. 77) have already been mentioned. Zo-

dariide is a large and widely distributed family which will probably be

found in the West Indies. Senoculide are running spiders found only

on the American mainland from Mexico throughout most of South Amer-

ica. The Archeide are interesting. There are three species of the type

genus, Archea, known from Baltic Amber but only one living species.

This species is known only from Madagascar. The only other genus in

the family is known by two species from Patagonia. The family Pla-

toride has also an interesting distribution,—two genera in South Amer-

ica and one in China. The remaining six families (Zoropside, Psech-

ride, Eréside, Prodidomide, Zodariide and Hersiliide) have a wider

distribution in the Old World.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 109

Of the Antillean families, all are found from north of Mexico to South

America except Palpimanide, which in America is known only from

South America, St. Vincent and Cuba, and Caponiide, which in America

is known only from Brazil to Mexico, St. Vincent, Hispaniola and Cuba

(elsewhere only in South Africa). Table I summarizes the Antillean

distribution of those families which are known from the Greater Antilles.

TasBLeE J.—Antillean distribution of families of spiders

: | Dy = | E S 2

ee a a a oa = 3

Sesh oe al hanya. | Se 5 s

| VASO 10 © ere x x x .- a ae x

Pileiaridee . 2... 2 ses x . <x x x x a

_ LEE . . . ie - x se

MAT ps a hn ew oe . . x a vas

1 ICE be re >< S< = x x

~ Lig Cr x me x x x e

Meetipidee- i.e ess x x Sc 4

Myetervd. 52... ss. 4 ie x ar 4

Beem ek >< ae vi 4 Sé

2 0 x al 2s . x

Balpamanide..:....-.... Le OS “s ate 2 x

WON sk x x x x 4 x

WWeridudse....)........ < ySilangd aS x x Si

Premppniide. ke. gee adil y yas x x A

Pee Sw. < x Sed Ne x x x

J) EG a . ae x be on s -

MMOMNISIO®...-. .... x ae x x x x Be

Brmbennide . se. ye x > eRlls Migiows a x x x

22 5, LG i .- Dee ait eats ip x Ee

Lr -- x SE be! bs x 4 x

eS es ~< x Si he te x Be

Le x x Sr) Me x x 4

21 LS 2S re 4 We SGie abs AS 4 4 x

It should be remembered that throughout this paper Lesser Antilles

means little else than St. Vincent, since it is the only one of these islands

which has been carefully studied. The need of work in Hispaniola is

emphasized by the fact that one man collecting for only a few days on

the small islands of Mona and Desecheo found representatives of twelve

families, while but thirteen families are recorded from the large neigh-

boring island of Hispaniola. It is strange that but nine families are

known from Jamaica. This island has had the benefit of several workers,

but poverty of its fauna is shown in other groups, so that this small num-

ber, while certainly not representing all the families to be found there,

may be significant.

110 ANNALS NEW YORK ACADEMY OF SCIENCES

In view of the careful work done on St. Vincent, it is not likely that

its number of families will be greatly increased. It lacks Dictynidex, but

this family is in the Greater Antillean list only by reason of a specimen

from Guane, Cuba. It lacks Gicobiide, of which we found a species on

Mona, but the family is otherwise unknown in the Antilles. Simon be-

heves the species we found on Mona to be carried by commerce. It is

curious that it should be on Mona, where there is almost no commerce, and

not on St. Vincent, where there is a great deal. Mimetide are now known

from Cuba and Porto Rico but not St. Vincent. They do not seem to be

common even on the first mentioned islands as they had not previously

been recorded and we have but one locality for each island. Finally,

Pisauride, although not common, are recorded from all the Greater An-

tilles and from Vieques but from none of the Lesser. As they are, for

the most part, rather large spiders, this is probably not due to defective

collecting. It is probably one of those defects in distribution which are

likely, when properly understood, to give us most valuable clues to the

larger problems involved.

DISTRIBUTION OF GENERA

As a rule, family is somewhat too large a group to be of much value in

a discussion of distribution and species is too small and uncertain, being

not only no less human than other taxonomic groups but even more

likely to be based on geographic range—the very thing we would like to

use it to study. Accordingly genus seems the most favorable group to

consider intensively.

The two Cohors of Petrunkevitch—Araneus and Dendryphantes—will

be left out of consideration and also the following genera: Ischnocolus,

Mygale, Blechroscelis, Teutana, Agobardus, Pholcus and Hasarius. The

first two are not considered good genera; the next three are recorded from

“West Indies” without further data; and the last two are said to be found

throughout the tropical countries but no West Indian records are known.

LESSER ANTILLES

There are 108 genera of which 54 or just 50 per cent. are not known

from the Greater Antilles. Seven, or 6.5 per cent., are peculiar to the

Lesser Antilles. Ischnothyreus is not known elsewhere in America but

is found in western Africa, Ceylon and the Philippines. Likewise, Corin-

nomma is known elsewhere only in Africa, eastern tropical Asia, Ceylon,

Malasia, Papuasia, Philippines and Australia. In addition to the seven

mentioned above there are 40 others which are not found in the Old

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 111

World. In other words, 43.5 per cent. are strictly American. Of these,

one, Alcimosphenus, is known from Greater Antilles but not from the

continent, 33 are known from South America, 21 are known from Cen-

tral America, 14 are known from United States.

There are 59 genera which are known in America outside of the Lesser

Antilles and are also known from the Old World. One of these, Dry-

musa, 1s known only from St. Vincent and Mona in the New World. In

the Old World it is known only from Cape of Good ae one species

being found in each hemisphere. Of the other 58, 56 (17) are known

from South America, 44 (6) are known from Central America, 41 (4)

ave known from United States. Leaving out of account cosmopolitan or

at least cosmotropical genera and those which are distributed pretty gen-

erally throughout the American-mainland, we have the numbers shown in

parentheses. Let us now examine the distribution of these more in detail.

One of them, Theone, has one species in St. Vincent, one in Missouri,

and the genus is represented also in France and Sumatra. Hahnia has

several species from Virginia and Wyoming northward to Greenland, two

in Patagonia and Terra del Fuego and one in St. Vincent and Porto

Rico. It is unknown from the middle of the western mainland. In the

eastern hemisphere it has been found in Europe, in the mountains of

India, in Japan and Sumatra. Bolostromus is found in Alabama (one

species), northwestern South America (four species), St. Vincent (one

species) and southwestern Africa. Oxzyopeidon has six species in Central

America and Mexico, one of them reaching to Arizona, one species in

St. Vincent and Mona, and in the Old World it is found in tropical east

Africa, India and Indo-China.

One genus found in Central America has just been ait! ‘Phe.

remaining five are found in both Central and South America but are not

known from United States. One, Beata, is known only from this section

of tropical American mainland, St. Thomas, St. Vincent and Africa.

The details of the distribution of Miagrammopes, Dysderina, Artema and

Stephanopis may be had by consulting the taxonomic part of this paper.

One is found in the Mediterranean region, three in Africa, three in Mada-

gascar, two in continental Asia, one in the Philippines, two in Malasia,

one in Melanesia, two in Polynesia and two in Australia.

Seven of the genera under discussion which are found in South Amer-

ica have been mentioned. The followimg ten are not known from the

American mainland north of Panama and, with the exception of Opopea,

which we found on Desecheo, not even from any of the Antilles except

St. Vincent: Accola, Theotima, Oonops, Opopea, Janulus, Episinopsis,

Dyschiriognatha, Ogulnius, Anapis, Caloctenus. One is found in Europe,

112 ANNALS NEW YORK ACADEMY OF SCIENCES

five in Africa, one in eastern Asia, two in Malay peninsula (one not being

known elsewhere in the Old World), four in Ceylon (one not being known

elsewhere in the Old World), one in New Caledonia, one in Japan, three

in the Philippines (two not being known elsewhere in the Old World),

two in Malasia and one in Australia.

I believe it is impossible to explain these distributions by accidental

dispersal in relatively recent times through wind or otherwise or by any

recent system of land bridges.

Taking into account all the genera found in the Lesser Antilles, we

find that 22 (20.4 per cent. of the total) are practically world-wide in

their distribution. This leaves 39 genera which have a more or less re-

stricted range in the eastern hemisphere. Of these, 7 (18.0 per cent.)

are found in central or northern Europe, 10 (25.6 per cent.) m the Med-

iterranean region, 23 (59.0 per cent.) im Africa, 5 (12.8 per cent.) m

Madagascar, 20 (51.3 per cent.) in continental Asia, 10 (25.6 per cent.)

in Ceylon, 8 (20.5 per cent.) in Japan, 8 (20.5 per cent.) in Philippines,

12 (30.8 per cent.) in Malasia, 5 (12.8 per cent.) in the smaller Pacific

Islands, and 8 (20.5 per cent.) in Australia.

The American distribution outside of the Lesser Antilles of Lesser

Antillean genera is shown in Table II. The study of this table had better

be deferred until after the Greater Antilles have been considered.

Taslte I].—American distribution of Lesser Antillean genera of spiders*

Not |

on | SAL) (A OS. 8 As) eae a -

main- only only only CA. | U.S. | U.S. |) 283 eee

land |

Not in other An-

pti |= eee Spey eae ee 9 19 | # 1 6 2 1 14 5+

Bo Henly 5:5 1 1 2 1 1 7

Hisp. only. ..... 1 1 2

Cuba only...... 3 i 2 1 3 10-

Jamm..; qubyn.3..'.' 1 1

Po wap. 2 32 1 L

P. R., Cuba a age ae 1 as 3 +

Hisp., Cuba.... es = = <e 7 ae ‘ 3 4

Hisp.. Jam..... Bie mp ue a ze a ee 1

P: +k ire pe

Calazc. 35-45% 3 9 12

P.. oR) eae

Jam. 34 ee. 3 De a a2 si 2 z

Hisp., Cuba, Jam. 1 = = rp 1 2

PL ES] Bass.

Cuba, Jam... we 1 1 Es 1 ss oF 5 8

Total. 11 25 2 3 16 5 3 43 108

* Columns refer to the mainland and rows to the Antilles. “C. A.” includes Mexico,

and “U.S.” includes everything north of Mexico.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 113

Porto Rico

Politically Porto Rico includes Culebra, Vieques, Desecheo and Mona.

In Table II and in what follows these islands are included when Porto

Rico is mentioned unless the context clearly implies a distinction. It

seems that the only error this might cause would be to make the con-

clusion concerning the cosmopolitan character of the Porto Rican fauna

less clear cut than it would be if we considered only the main island.

There are 51 genera known from the Porto Rican islands of which

cnly 5 or 9.8 per cent. are not known from the other Antilles. It is

interesting that four (Stichoplastus, Gicobius, Mecolesthus and Edricus)

of these five are added to the fauna by this list, while the other one

(Pardosa) is credited as peculiar in the Antilles merely because a dubious

taxonomic shift has taken a Cuban species out of the genus. No known

genera are peculiar to the Porto Rican islands. All of these five genera

are known elsewhere in America; one being found elsewhere only in

South America; two only from South America to Mexico; one from

Brazil to Massachusetts, in Azores, Canaries, Mediterranean region,

Arabia, Japan and New Caledonia; and the fifth is practically cosmo-

politan.

Unfortunately the standing of Prostheclina is in considerable doubt.

As Simon considers it in part a synonym of Sattis it would be difficult

to determine its Old World distribution. For details see page 107. It

will be omitted from further discussion. As it occurs in the other An-

tilles, whether it be a synonym of Satis or not, this omission does not

affect the statements of the preceding paragraph.

Of the 50 remaining genera, 15 (30 per cent.) are restricted to Amer-

ica (including Wala, which is found in Bermuda). All of these are

known from the continent, being distributed as follows: 12 are known

from South America, 12 are known from Central America, 9 are known

trom United States.

Of the 35 genera found also in the Old World, only one, Drymusa, is

not known from the American mainland. As has been stated its distri-

bution is Mona, St. Vincent and Cape of Good Hope. Of the remaining

34, 33 (5) are known from South America, 32 (4) are known from

Central America, 28 (2) are known from United States.

Leaving out, as was done in considering the Lesser Antilles, the rather

generally distributed genera, we have the numbers shown in parentheses.

They refer to Miagrammopes, Dysderina, Opopea, Artema, Hahnia, and

Oxyopeidon. The details of their distribution may be had by reference

to the taxonomic part of this paper. All are found in the Lesser Antilles

and only one (Jiagrammopes) elsewhere in the Antilles.

ze ANNALS NEW YORK ACADEMY OF SCIENCES

Taking into account all the genera except Prostiteciina known from

the Porto Rican islands, we find that 22 (44 per cent. of the total)

are practically world wide in their distribution. ‘This leaves 15 genera

which have a more or less restricted range in the eastern hemisphere.

Of these, 1 (7.7 per cent.) is found in central Europe, 6 (46.2 per cent.)

in the Mediterranean region, 11 (84.6 per cent.) in Africa, 3 (23.1 per

cent.) in Madagascar, 9 (69.2 per cent.) in continental Asia, 1 (7.7 per

cent.) in Ceylon, 3 (23.1 per cent.) in Japan, 3 (23.1 per cent.) in Philip-

pines, 4 (30.8 per cent.) in Malasia, 2 (15.4 per cent.) in the smaller

Pacific islands and 5 (38.5 per cent.) in Australia and New Zealand.

The American distribution outside of Porto Rico of Porto Rican

genera is shown in Table III. It is worth noting in passing that of the

Tapte II].—American distribution of Porto Rican genera of spiders?

Not | S. A.

on | SeAg (|) C. Al) UL S20 oS. ARG. cA S32 Ae eee

main-| only | only | only | C.A. | U.S. | U.S. | U.S. | Total

land |

Not in other An-

Gilles Mase ecier. an 1 2 2 5

Lesser only..... 1 i 2 if 1 it 7

Iaiisyo, “Olly. 54 5 - 1 bh Se il 2

Cuba onliyncoeee if 2 3

Less., Hisp..... ses 2s Ve oe 1 uy

hess. “Cuba.. -or a. a a 1 3 -

Cuban Janie ne aR on He ne ae if 1

Less., Hisp., Cuba Sul, Aes He a De 9 12

Less., Cuba, Jam. | 2 2

Hisp., Cuba, Jam. 5 5

Lesser, Hisp.,

Cuba, diame = Ec ik a, 1 6 8

OG ale eer 1 3 1 1 8 2 1 33 50

50 genera considered, 34 are in the Lesser Antilles and 38 in the Greater.

This would show a slightly greater affinity of the Porto Rican fauna with

the islands to the west. However, many of these are widely distributed

genera. It was seen above that all of the five genera having more or

less restricted range and found also in the Old World are found in the

Lesser Antilles and only one in the islands to the west. Adding Phor-

mictopus, Hugnatha, Wulfila, Aysha, and Stoidis—the only strictly Amer-

ican genera found in Porto Rico which are not wide-ranging on the

mainland—we find 8 in the Lesser Antilles and 6 in the islands to the

west. It must be concluded that, as far as genera of spiders are con-

’ Columns refer to the mainland and rows to the Antilles. ‘‘C. A.” includes Mexico,

and “U.S.” everything north of Mexico.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 115

cerned, the Porto Rican fauna is as closely related to the Lesser Antillean

as it is to that of the other Greater Antilles. This conclusion is strength-

ened by remembering that St. Vincent, an island well down in the string

of Lesser Antilles, furnished almost all the data for that group of islands.

HISPANIOLA

Although some of the records for this island are credited to the Haitian

portion of it through the unfortunate use of “Haiti” for the whole island,

nevertheless most of them really came from Haiti, especially near Port

au Prince, being those of Banks’s list. It seems best to consider the

island as a whole until we know more about the differences between the

parts.

There are only 52 genera known from Hispaniola. This, clearly, is

but a small part of its fauna, and since the more interesting genera, those

of restricted range, are probably most abundant in the almost totally

unstudied mountains, it is possibly not a fair sample. However, 9 genera,

or 17.3 per cent., are not known elsewhere in the Antilles and 2 (Scope-

lobates and Bythocratus), or 3.8 per cent., are peculiar to the island.

Trichopelma is known elsewhere only from South America: Tobias only

from Central America and northern South America; Acacesia only from

southern United States to Panama (a single species throughout) ; Weta-

cyrba, the same distribution except that it is also found in Buen Ayre,

off the northern coast of South America; Jcius, especially in United

States (as far as American records go), Mexico, Uruguay, Europe, Africa

and Asia: Linyphia and Cyrtophora, practically cosmopolitan but not

recorded from the other Antilles.

Again leaving out Prostheclina, we find 24, or 47.1 per cent., of the

remaining 51 are strictly American (including Wala and Futichurus,

which are known from Bermuda). Of these Alcimosphenus is known

from St. Vincent, Cuba and Jamaica, and Nilacantha from Jamaica, but

neither from the continent. These, with the two peculiar genera, leave

20 which are distributed as follows: 16 are known from South America,

17 are known from Central America, 11 are known from United States.

There are 27 Hispaniolan genera known elsewhere in America and

also in the Old World. All are found in continental America and as

follows: 27 (2) are known from South America, 25 (1) are known from

Central America, 24 (1) are known from United States.

As before, the numbers in parentheses refer to genera of more restricted

range. As a matter of fact only two are concerned: Wiagrammopes,

known from all the Greater Antilles except Jamaica, from St. Vincent,

Brazil to Mexico, Africa, Madagascar, southern Asia and Australia: and

116 ANNALS NEW YORK ACADEMY OF SCIENCES

Syrisca, known from Cuba, southwestern United States, Brazil, Paraguay

and Africa. ,

Considering all Hispaniolan genera except Prostheclina, 21 or 41.2

per cent. are almost world wide in their distribution. This leaves six

(IMiagrammopes, Anyphena, Syrisca, Trachelas, Corinna and Icius) to

be considered. Of these 1 (16.7 per cent.) is found in Europe, 2 (33.3

per cent.) in the Mediterranean region, 5 (83.3 per cent.) in Africa, 2

(33.3 per cent.) in Madagascar, 5 (83.3 per cent.) in continental Asia,

i (16.7 per cent.) in Japan, 2 (33.3 per cent.) in Malasia and 1 (16.7

per cent.) in Australia.

The American distribution of Hispaniolan genera outside Hispaniola

is shown in Table LV.

Taste 1V.—American distribution of Hispaniolan genera of spiders®

Not S.A

on S. A CO. ts S. As Cons Sp Ci

main-| only only only CA 4) US a suis Us Total

land

Not in other An-

Pulleses acer 2 a 1 1 t a 9

Lesser only..... 1 1 2

Pr rk: onlye sac. 1 i 2

Cuba only. 42~ 7 1 il 1 1 +

Jam only. 25: 1 ze = a ce es a Re) 1

hess, Pe eRe’. 15% ay = oe ae ae Re sg 1 Lo

Less., Cuba.) .. ve af ae a if Se 3 40

Less... Jams oj: se a fe oN Ae ] am _- 1

Less., P.R., Cuba. he oe we be 5) ¥. 4 9 12

Less., Cuba, Jam. it 1 ; 2

PWS tite ae ana.

A ee ee if 5 6

Less., P. R., Cuba,

JMS cae eres ik 1 6 8

Totaly ci. 3) 3 2 0 8 2 2 30 52

CUBA

There are 82 genera recorded if Misumessus echinatus Banks be put

under Misumenops and Dolomedes (see Thaumasia) be omitted. Of

these, 24, or 29.3 per cent., are not known from the other Antilles, but

only one (Hapalopinus), or 1.2 per cent., is peculiar to Cuba. Of the re-

maining 23, all are found in continental America. Oxysoma is confined

to South America except for a single Cuban species. Filica and Thana-

tidius are American genera and have but two species each on the main-

° Columns refer to the mainland and rows to the Antilles. ‘C. A.” includes Mexico,

and “U.S.” everything north of Mexico. :

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 117

land, and in both cases one is in southeastern United States and the other

in Brazil, but the Cuban species of Hilica is distinct, while that of Tan-

atidwus is common to Cuba and United States. Drezvelia is found through-

out most of the world’s tropics and subtropics, including Central America

and United States, but has not been reported from South America or any

of the Antilles except Cuba, while Peckhamia is known only from Pan-

ama northward on the American mainland and in Cuba. Callilepis,

Mysmena, Ceratinella and Hyctia are known in America only from Cuba

and the mainland north of Mexico, but they are all found in Europe and

elsewhere in the northern hemisphere. ‘The other 14 are all widely dis-

tributed in America; Hurypelma, Ceratinopsis, Cupiennius and Thiodina

being strictly American; Smeringopus, Inthyphantes, Trbellus, Tegenaria

and Myrmarachne being at least cosmotropical: Dictyna occurring in

Europe and Asia, including Japan and the Philippines; Theridula, in

the Mediterranean region, Africa, Madagascar, tropical and eastern Asia,

Japan and the Philippines; Bathyphantes, in Europe, Asia, New Zealand

and Australia; Gea, in western Africa, tropical Asia, the East Indies,

Polynesia and Australia; and Mangora, in the Atlantic islands, Europe,

Asia and Ceylon.

Prostheclina is troublesome, as before, and will be omitted. , Including

Hutichurus and Wala, which are found in Bermuda, 31, or 38.3 per cent.,

are restricted to America. With the exception of Hapalopinus, which is

confined to Cuba, and Alcumosphenus, which is found in Jamaica, His-

paniola and the Lesser Antilles, all these are found on the mainland and

are distributed as follows: 22 are known from South America, 20 are

known from Central America, 18 are known from United States.

All of the 50 Cuban genera which are found in the Old World are also

found on the American mainland. They are distributed as follows: 44

(3) are known from South America, 45 (3) are known from Central

America, 46 (6) are known from United States.

As before, the numbers in parentheses refer to genera having a more or

less restricted range. MJiagrammopes is found from Brazil to Mexico, in

St. Vincent and all the Greater Antilles except Jamaica, in Africa, Mada-

gascar, southern Asia and Australia. Callilepis is widely distributed in

the Old World but is confined to Cuba and north of Mexico in America.

Mysmena is found in Cuba, southeastern United States, France, northern

Africa, Ceylon and the Philippines. Ceratinella is known from Cuba,

northeastern United States to Labrador and northern Eurasia. Stepha-

nopsis ranges from Patagonia to Panama, in Tortola (Lesser Antilles),

Cuba, Madagascar, Malasia, Melanesia, Polynesia and Australia. Syrisca

is recorded from Brazil, Paraguay, southwestern United States, His-

118 ANNALS NEW YORK ACADEMY OF SCIENCES

paniola, Cuba and Africa. Hyctia is in Cuba, most of the United States

and Europe. Pellenes occurs in Cuba; Jamaica, Central America and

northward, Europe, Mediterranean region and northeastern Asia.

Taking into account all the Cuban genera, we find that 29 (35.8 per

cent. of the total) are practically world wide in their distribution. Of

the remaining 21 which have a more or less restricted range in the Old

World, 7 (33.3 per cent.) are found in central and northern Europe; 9

(42.9 per cent.) im the Mediterranean region: 13 (61.9 per cent.) in

Africa; 5 (23.8 per cent.) in Madagascar; 15 (71.4 per cent.) im conti-

nental Asia; 2 (9.5 per cent.) in Ceylon; 4 (19.0 per cent.) in Japan;

4 (19.0 per cent.) in the Philippmes: 4 (19.0 per cent.) in Malasia; 3

(14.5 per cent.) in the smaller Pacific islands and 8 (38.1 per cent.) in

Australia and New Zealand.

The American distribution of Cuban genera is shown in Table V.

TABLE V.—American distribution of Cuban genera of spiders *

Not | | S. A.

on, | S.A: | CC. A. 4 DLS) S.A. | CA See

main-| only only only C.A. | U.S. | U2 SOs See

land

eee aa a | Sn | eee

Not in othet An-

tales ois Si i iT - Se 2 2 14 24

Lesser only.... 3 x 2 1 3 10

PP gaye s c= - ca 2 3

Hisp. only,..-.. ie mn 1 ; 1 : 1 1 +

Jam: iuanily. o> 5c. Bt ee. rs = =e Z ai 2

Leas. Pele 1 zi 3 -

Tess... Ehispe 2 24). 1 3 -

PL hi. Jam | | me 1 1

Less., P_E., Hisp- | | 3 9 12

Less., P. B., Jam. VAs nh 2 2

Less., Hisp., Jam. © 19) if | ; 2

P.R.,Hisp, Jam. | .. | | | 5 5

Less., P.R., Hisp., | |

Page Bok oe Ah 1 1 6 8

ital e 5. es RE BS 6 9 5 4 | 49°) ‘Bf

JAMAICA

Only 31 genera of spiders are known from Jamaica, including Pros-

theclina. Of these 7, or 22.6 per cent., are not found in the other An-

tilles. The question as to whether Prostheclina of Keyserling should be

treated as a synonym of Saitis of Simon has been mentioned several times

and Prostheclina has been omitted from the discussions when world-wide

70 Columns refer to the mainland and rows to the Antilles. ‘“‘C. A.” includes Mexico,

and “U.S.” everything north of Mexico.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 119

distribution was concerned. In Jamaica we are troubled by the fact that

three species of Saitis are recorded. There are three other species in

South America and the genus in the broad sense is known from south-

western Europe, western and southern Africa, India, Malasia, Polynesia

and Australia. If we include the records for Prostheclina with Sastis

the result will be to decrease the apparent distinctness of Jamaica from

the other Antilles, but as it is still quite distinct this would probably be

the fairest course.

We then have 30 genera credited to Jamaica of which 6, or 20 per cent.,

are not known from the other Antilles. Of these 3, or 10 per cent., are

‘not known elsewhere in America. Theridionexus and Epecthinula are

confined to Jamaica and Cyatholipus has been found elsewhere only in

South Africa. Fourteen genera (43.3 per cent.) are strictly American ;

10, or 33.3 per cent., are practically cosmopolitan or cosmotropical. In

addition to these and Sastis and Cyatholipus, which have just been met-

tioned, Pachylomerus is known from Jamaica, St. Vincent, Brazil (?),

Venezuela to Maryland, the Mediterranean region and Japan. Ischno-

thele is found in Jamaica, South America to Mexico, Bahamas and either

it or a closely related genus, which we will consider as it, in eastern

Africa, Madagascar and India. Filistata is known from South America,

including the Galapagos, to California and southeastern United States,

Bermuda and the Atlantic islands, Mediterranean region, Africa, central

Asia, Philippines and Australia. Pellenes is reported from Jamaica,

Cuba, Central America to Canada, Europe, Mediterranean region and

northeastern Asia. The numbers are so few that the reader can easily

analyze the distributions from the data just given.

Of the 10 American genera which have been found on the mainland,

6 are known from South America, 10 are known from Central America

and Mexico, 7 are known from United States.

The American distribution of Jamaican genera is shown in Table VI.

120 ANNALS NEW YORK ACADEMY OF SCIENCES

TaBLeE VI.—American distribution of Jamaican genera of spiders™

Not S. A

on Se Ae Ce U.S S. A WA. 12s. As. eee

main-| only only only. | CA. | U. S._|-U. 8. | USSetene

land

Not in other An-

tillese = 3 ae 3 at 1 oe cL 1 6

lesser only.2--- ee 1 it

Hisp: -onlysa2e 1 I

Cuba only. 4-2. : 2 2

ess. akuispee wake Bek Se oy ate ay 1 ie eee is 1

PR; Cubase. = Bt ae bie ae a ee if 1

Less., P.R., Cuba a ee ig se a — 2 2

Less., Hisp.,Cuba 1 - se ie if 3 ee x 2

P. R., Hisp., Cuba Bis 1 oh a af = sR ) 6

Less., P. R., Hisp.,

Guibas. cere se ne iJ Wee i ie a4 6 8

Totals pcre 5 at 2 0 3 3 0 16 30

GREATER ANTILLES

Omitting the two cohors and the seven doubtful genera mentioned on

page 000; also considering that Misumessus echinatus is a Misumenops.

that the Antillean records for Prostheclina in Petrunkevitch’s Catalogue

should be credited to Saitis and that Dolomedes has not been recorded

from Cuba, there are 117 genera left, of which 63, or 53.8 per cent., are

not known from the Lesser Antilles. Six, or 5.1 per cent., are peculiar

to the Greater Antilles. They are Hapalopinus in Cuba, Scopelobates in

Hispaniola, Theridionexus and Hpecthinula in Jamaica, Bythocrotus in

Hispaniola and Nilacantha in Hispaniola and Jamaica. Cyatholipus is

not known in America outside of Jamaica but occurs in South Africa.

in addition to the six mentioned above there are forty-four others which

are not found in the Old World. In other words, 42.7 per cent. are

strictly American. One of these, Alcimosphenus, is also known from the

Lesser Antilles (St. Vincent) but not from the mainland; 33 are known

from South America, 30 are known from Central America and Mexico,

24 are known from United States.

There are 66 genera which are known in America outside of the

Greater Antilles and are also known from the Old World. One of these,

Drymusa, is known only from Mona, St. Vincent and Cape of Good Hope.

Of the other 65, 58 (9) are known from South America, 56 (7) are

1 Columns refer to the mainland and rows to the Antilles. ‘“‘C. A.” includes Mexico,

and “U.S.” everything north of Mexico.

~~, A

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 121

known from Central America and Mexico, 55 (7) are known from United

States.

As before, the numbers in parentheses refer to genera of more or less

restricted distribution. J/ysmena, Ceratinella and Hyctia are Holarctic

genera which are for the first time in West Indian list and now only by

reason of being found in Cuba. The others (/schnothele. Miagram-

mopes, Dysderina, Opopea, Artema, Stephanopsis, Syrisca, Hahnia, Oxyo-

peidon, Pellenes and Saitis) are considered rather fully in the discussions

concerning the individual islands and also in the taxonomic part of this

paper. |

Of the 117 genera in the Greater Antilles, 34 (29.1 per cent. of the

iotal) are practically world wide in their distribution. This leaves 33

genera which have a more or less restricted range in the eastern hemi-

sphere. Of these 10 (30.3 per cent.) are found in central or northern

Europe; 13 (39.4 per cent.) in the Mediterranean region: 22 (66.7 per

cent.) in Africa; 7 (21.2 per cent.) in Madagascar; 23 (69.7 per cent.) —

in continental Asia; 3 (9.1 per cent.) in Ceylon; 7 (21.2 per cent.) in

Japan; 6 (18.2 per cent.) in the Philippines; 8 (24.2 per cent.) in

Malasia; 5 (15.2 per cent.) in the smaller Pacific islands and 9 (27.3

per cent.) in Australia and New Zealand.

The American distribution of genera found in the Greater Antilles is

shown in Table VII and those found in the Lesser Antilles are added

for comparison.

TABLE VII—American distribution of Antillean genera of spiders ®

Greater Totals

Greater and Lesser ps

only Lesser only Greater Lesser W. I.

Not on mainland... rs 2 9 9 af 18

Pee Only.....2... 5 5 19 10 24 29

2 2 af if 3 2 =

S800 ~ 2 1 6 3 7

oo 6 10 6 16 16 22

o 2 6 2 at 8 3 9

- 2 ee 4 2 3 6 5 9

ooo. A. U.S... 29 30 14 59 44 yes

SS es Sa ee ——— ee SS

_ ere 63 54 54 117 108 EYAL:

2 «“C. A.” includes Mexico, and “U.S.” everything north of Mexico.

122 ANNALS NEW YORK ACADEMY OF SCIENCES

DISTRIBUTION OF SPECIES

Table VIII summarizes the data at hand. A few doubtful species

have been omitted and several undescribed ones have been included.

TaBLeE VIII.—American distribution of Antillean species of spiders *®

Not S.A

on S. A Ch: U.S S. A C. Ac Sa Cie

main- only only only CUA We. es U.S Total

land

Lesser only..... 88 18 2 2 4 2 di ~ 121

P. Be enkys 2 22 6 2 : aye 2 1 1 12

Hasp: only. = 3:7 18 if 1 2 1 4 1 28

Cuba Uonlvese.-- 33 2 3 22 se § i 5 74

Jaime, Orby. oo. 23 ] 1 1 26

Less PR oe 9 1 1 bs 11

Less.,-Hispooc: 1 ae 1 2

Bess: “Cuban a 3 ee 2 i 2 8

PRS isp. es 2 2 a 2 oe 7

PL i Cabae 2 2 1 2 2 9

Hisp. Cuba: ot: 5 2 3 10

Easp., Jam eee, 5 5

Guba, Jam zc... 1 ms 1

Less., P.R., Hisp. 2 1 es 3

Less., P.R., Cuba 2 1 i 2 6

Less.. Hisp., Jam. 1 1

P:R” (rash.

Saat es ae “e 1 1

Hisp., Cuba, Jam. il i

Less., P.R., Hisp.,

Gubay see 2 af 2 2 2 9

Less., P. R., Cuba,

a Ih nen os ee ss 2 oe ae = 2 2

eY)

ate 2. Eee ee i ge ee yes 2 md 2

All (2) Antilles. 1 3 4

Meped. oe 206 |" 2a) a0 A aoD red a emo 6 | 30 | 347

GENERAL DISCUSSION

Is THE FAUNA OF THE LESSER ANTILLES DISTINCT FROM THAT OF THE

GREATER ANTILLES ?

Comparing the two usual divisions of the Antilles—Greater and

Lesser—we see that 50 per cent. of the Lesser Antillean genera of spiders,

unfortunately meaning little more than the fauna of St. Vincent, have

78 Columns refer to the mainland and rows to the Antilles. “C. A.” includes Mexico,

and “‘U. S.”’ everything north of Mexico.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 123

rot been found in the Greater Antilles and 53.8 per cent. of those known

from the Greater Antilles have not been reported from the Lesser An-

tilles. In the Lesser Antilles 6.5 per cent. of the genera are peculiar

and in the Greater, 5.1 per cent. In the Lesser Antilles 43.5 per cent.

are strictly American and in the Greater, 42.7 per cent. The last two

comparisons indicate considerable similarity in the general character of

the two faunez and raise doubt as to the significance of the fact that half

of the fauna of each division is distinct from that of the other division.

Do the two divisions have significantly distinct faune? Taking His-

paniola because it is in the midst of the Greater Antilles and bearing in

mind that further study of the Hispaniolan spiders is more likely to

bring out differences than similarities because we are now ignorant of

the fauna of the interior, we note that 42.3 per cent. of its genera are

not known from the Lesser Antilles, 44.3 per cent. are not known from

Porto Rico, 30.8 per cent. are not known from Cuba and 65.4 per cent.

are not known from Jamaica. The case of Jamaica is probably due, in

large part, to ignorance and a further study of Cuba and Porto Rico

without a further study of Hispaniola would doubtless reduce their fig-

ures, but it must also be kept in mind that data for the Lesser Antilles

is largely drawn from an island well down the line, so that we must con-

clude that with respect to the genera of spiders the Lesser Antilles are no

more different from the Greater Antilles than the different islands of

the latter are from each other.

Of course, species show a much more limited geographic range than

genera and the actual distance of St. Vincent from the Greater Antilles

apart from purely faunistic factors becomes an important factor when

considering species. ‘There are 168 species known from the Lesser An-

tilles of which 72.0 per cent. are not reported from the Greater Antilles,

and of the 226 species known from the Greater Antilles 79.2 per cent.

are not known from the Lesser. In the Lesser Antilles 52.4 per cent.

of the species are not known elsewhere and in the Greater Antilles, 42.5

per cent. It will thus be seen that the spider fauna of the rather isolated

Lesser Antilles is not much, if any, more specialized than that of the

Greater. Again taking Hispaniola for a basis of comparison we find

that of its 76 species, 73.7 per cent. are not known from the Lesser

Antilles, 68.4 per cent. are not known from the Porto Rican islands,

60.5 per cent. are not known from Cuba and 77.6 per cent. are not known

from Jamaica. To say the least, the material at hand would not justify

us in considering the Lesser Antilles any more distinct from the Greater

than any of the islands of the Greater are from each other.

124 ANNALS NEW YORK ACADEMY OF SCIENCES

Is THE ANTILLEAN FAUNA DISTINCT FROM THAT OF THE MAINLAND?

The introductory paragraph to Scharff’s Chapter on the origin of the

West Indian fauna is as follows:

“North and South America are to be regarded. according to Professor Suess,

as two essentially distinct land-masses, between which is interposed, as a third

element, the area of Central America and the Antilles. This geological dis-

tinctness of Central America and the Antilles from the two neighboring conii-

nents is scarcely recognizable in the fauna of the great isthmus. But the

West Indies are comparable to a wedge driven in between two faunistically,

more or less, independent and distinct masses. Almost everyone who has dealt

with the fauna or flora of the West Indian islands has expressed his surprise

at this fact. In position. says Dr. Wallace, the Antilles form an unbroken

chain uniting North and South America, in a line parallel to the great Central

American isthmus. Yet instead of exhibiting an intermixture of the produc-

tions of Florida and Venezuela, they differ widely from both these countries,

possessing in some groups a degree of speciality only to be found elsewhere in

islands far removed from any continent.”

Several years ago (1913) I published a brief note on the distribution

of occidental spiders, getting the data entirely from Petrunkeyvitch’s

catalogue. As has been done here, I roughly divided the western hemi-

sphere into four parts and found that 59.8 per cent. of the South Ameri-

can genera of spiders were not known elsewhere in America, 21.5 per

cent. of those in Central America and Mexico were peculiar to that region

and 37.5 per cent. of those from United States and Canada were not

found farther south. Of the 133 Antillean genera then considered 13.6

per cent. were known only from the West Indies. The present paper

includes 171 genera without adding any peculiar ones, so that the per-

centage is reduced to 10.5. This would also reduce the percentage for

some of the other divisions but the figures as they stand are about as

reliable as the data on which the revised ones would be based, genus

being an indefinite sort of thing, so that we may conclude that the genera

of spiders do not confirm the idea of the distinctness of the West Indian

fauna. Instead of being very distinct from the mainland they are less

so than any of the mainland divisions are from each other. Is this be-

cause spiders are more generally distributed than other animals or is the

impression of distinctness an erroneous one based on a consideration of

special cases ?

The best way to settle the question of the distinctness of the Antillean

fauna is to study carefully a number of groups which are fairly well

represented in the Antilles and we are doing this in connection with the

survey of Porto Rico which is being carried on under the auspices of the

New York Academy of Sciences. Mammals were used by Wallace with

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 125

wonderful success but they do not seem to be good material for a West

Indian discussion because there are so few of them. It is decidedly in-

teresting that, except for bats, the Antillean mammals are rodents be-

longing to South American groups and Solenodon, a genus of Insectivora,

whose nearest relatives are found only in Madagascar. ‘These relation-

ships will be referred to later but it may be noted in passing that such

distributions are fairly common among spiders. ‘The figures given by

Wallace for birds show that about one-third of the Antillean genera are

peculiar. This proportion is far greater than that for spiders and, when

Fie. 3.—Sandy plain south of Pinar del Rio, Cuba

compared with the bird fauna of Central America, for example, shows a

high degree of speciality, but Wallace’s list gives only 95 genera of birds

for the West Indies and there are more than 500 genera in Central

America. The small number of vertebrates in the West Indies and the

fact that a large proportion of those which are there are peculiar seem

to be evidences of the unfitness of the Antillean environment for yverte-

brate life rather than of any special distinctness of the Antilles from the

standpoint of geographic distribution.

It seems almost pedantic to point out the necessity of considering en-

126 ANNALS NEW YORK ACADEMY OF SCIENCES

92Taphic distribution and vet it is almost

never taken up in detail. There are several very practical reasons for

eae :

ss a 7 bere Pg el ee ene ee

are Well liiustrated DV this paper. First. there 1s

Lenadis: cia a, See ie lee ae a 7. ee

aiready Such a Mass Of detall that 1t Can scarcely be nandied and. second,

py Lad pe) eee Sy Ca ee oe 2 en Ps >

the author is oiten, as 1m this case. ignorant of the ecological conditions

under which the species occur throughout their ranges. The importance

of ecological conditions may be illustrated in connection with what has

= = = . 7 s tT

just gone before as well as with what follows

1G. 4—View near Banos San Vincente. Cuba

The level area is rich agricultural land. and even the cliffs support a rather luxuriant

vegetation.

Mr. Leng and myself went to Pifar del Rio, Cuba, chiefly to study the

ecological distribution of insects. We collected spiders in a number of

environments, two of which are of interest here. South of the city is a

sandy plain which we studied fairly intensively between the nime and

thirteen kilometer posts on the road to Coloma. Leaving out of account

the really important local differences caused by ground water level and

the resulting differences in vegetation, figure 3 may be taken as fairly

typical of the region. North of the city are the mountains and we col-

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 127

lected especially in the vicinity of Bahos San Vincente. ‘The difference

in elevation between this place and the plains is not enough to cause a

noticeable difference in temperature apart from that connected with soil

conditions and moisture. Figure 4 gives an idea of the character of

the region—moist atmosphere, lime-humus soil, rather “tropical” vege"

tation. Leaving out of account the Cohors Araneus and Dendryphantes

for taxonomic reasons and not considering the minor ecological differ-

ences in each region, we found 10 genera on the plains which we did not

find at Banos, 24 at Banos which we did not find on the plains and only

9 genera at both places. That is, 52.6 per cent. of the plains genera

were not found in the hills and 72.7 per cent. of the hills genera were

not found on the plains. In other words there is at least as much differ-

ence between these two nearby but ecologically different localities as

there is between Cuba and the other islands or between the Antilles as a

whole and the mainland.

Furthermore, this must give us a pause in considering the relationships

of the various islands to each other and to the mainland, for 90 per cent.

of the genera we took on the plains are found in United States and only

77.3 per cent. of those from the hills; 40 per cent. of the former are

known elsewhere only in United States, as far as America is concerned,

and only 4.5 per cent. of the latter. The general impression is that the

West Indies are more closely related to Central and South America than

to United States and, going still further, that the Lesser Antilles are

especially related to South America and the Greater Antilles to Central

America. If this be true, in how far is it due to land bridges, ocean

currents or wind and in how far is it due merely to similarity, and hence

congeniality, of environment? ‘The first part of the question needs prior

consideration, and the second part cannot be satisfactorily answered until

we know more about animal ecology.

MAINLAND AFFINITIES OF THE ANTILLEAN FAUNA

As far as the data concerning the genera of spiders go, the Lesser

Antilles (St. Vincent) seems to have a closer affinity with South America

than does the other division of the Antilles, since 22.2 per cent. of the

genera of the Lesser Antilles are known elsewhere in this hemisphere

only in South America and the corresponding percentage for the Greater

Antilles is only 8.6. However, if we include all the genera whose dis-

tribution we can study we note that only 82.4 per cent. of the genera

known from the Lesser Antilles are known also in South America, while

the corresponding percentage for the Greater Antilles is 84.3 per cent.

The belief that work in the northern Lesser Antilles will still further

ANNALS NEW YORK ACADEMY OF SCIENCES

oa)

reduce the proportion of strictly South American genera in that division

makes us somewhat dubious of there being any marked difference between

the two divisions of the Antilles with respect to this relationship.

As far as the affinity between the Greater Antilles and Central America

(including Mexico) goes, we note that only 2.6 per cent. of the Greater

Antillean genera are confined, in their American mainland distribution,

to that region. Including all the genera, 73.5 per cent. of those known

from the Greater Antilles are known also from Central America and

Mexico, but the same percentage for the Lesser Antilles-is 60.2, which is

practically the same, in view of the fact that it is so largely based on

the St. Vincent fauna.

As was pointed out several paragraphs before, Cuba has a number of

genera known on the American mainland only from United States and

Canada, but such genera represent only 5.1 per cent. of the genera known

from the Greater Antilles (as a matter of fact, only 7.4 per cent. of the

Cuban genera), while the same percentage for the Lesser Antilles is 2.8.

Of the Greater Antillean genera 67.5 per cent. are known also in United

States and Canada and 50.9 per cent. of those reported from the Lesser

Antilles.

This, it seems to me, confirms the notion that, while individual genera

differ, the general make-up of the spider fauna of the Lesser Antilles

does not significantly differ from that of the Greater Antilles and that

neither division has drawn from, or given to, any particular portion of

the mainland much more than the other division.

A comparison of the Porto Rican spiders with those of the other

islands brings out some interesting points. It has, as far as we know,

no peculiar genera and only 9.8 per cent. of its genera are not known

from the other. Antilles. The data for the Lesser Antilles show 6.5 per

cent. of peculiar genera and 50 per cent. which are not known from the

other Antilles: for Hispaniola the figures are 5.8 per cent. and 17.3 per

cent.: for Cuba, 1.2 per cent. and 29.5 per cent.: and for Jamaica, 6.6

per cent. and 20 per cent. The fauna of Porto Rico seems, then, to be

of more recent origin than that of the other islands. This idea is

strengthened by the fact that, while the differences are not great, only

30 per cent. of Porto Rican genera are strictly American as compared

with 43.5 per cent. for the Lesser Antilles, 47.1 per cent. for Hispaniola,

38.5 per cent. for Cuba and 43.3 per cent. for Jamaica: also by the fol-

lowing, which is not strictly a restatement of the preceding, since a genus

may be found in the Old World without being so widely distributed as

to be classed as cosmopolitan. Cosmopolitan or cosmotropical genera

make up 44.0 per cent. of the Porto Rican genera as compared with 20

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 129

per cent. for the Lesser Antilles, 41.2 per cent. for Hispaniola (practi-

cally, the port towns), 35.8 per cent. for Cuba and 33.3 per cent. for

Jamaica.

Species also show the composite character of the Porto Rican spider

fauna and are perhaps more convincing than genera. Only 8.7 per cent.

of the Porto Rican species are peculiar as contrasted with 52.4 per cent.

for the Lesser Antilles, 23.7 per cent. for Hispaniola, 25.2 per cent. for

Cuba and 48.9 per cent. for Jamaica.

In view of this unusual character of Porto Rican spiders, it is worth

while discussing their distributicnal affinities. Table VIII gives the

deta for the American distribution of species in a condensed form.

Table IX shows the percentages of the total number of species in the

several islands for various groupings.

(ABH bx

Lif. PR: | Hisp. | Cuba. Jam.

phen s |

ee ee eee SEA. 52.4 Bia | 23.7 | Baa ARS

Mig eOiLHer ANGtIIEGS. 2.2. 62 ess tere oc | 1220 gO: a Wey oa | 56.5 Hots

In other Antilles but not on mainland. | 13.1 29.0 A276 ESE SBS:

On mainland but not in other Antilles. | 19.6 Sau A | ald 6:4

mantic ( tObal) 20.3... ke eae ess | 34.5 62.3 AB OF hd oh. 29.8

Porto Rico has as high a mainland affinity as Cuba and higher than

the other islands, but it has a lower direct affinity (a lower percentage

of species found on the mainland and not on the other Antilles) than

any except Jamaica. That is, its fauna is largely mainland species

which it has received by way of the other islands; Cuba’s species are also

largely mainland species, few of which have been passed on, while

Jamaica has few mainland species and most of these have been received

by way of the other islands or have been passed on to them. Of course,

it would be possible for a species to originate in a West Indian island

and then be transferred to the mainland and this may be true in Cuba.

It would be impossible to determine from the data whether this has hap-

pened or not but it probably did not occur often enough to seriously

complicate matters.

The rest of Table 1X agrees with the idea just stated and repeats what

was said before, as might be expected, since it is largely a complement of

the two lines just considered. Porto Rico has the smallest percentage of

endemic species of any of the islands or groups of islands; and, partly

for this reason, but partly also because it has received a large proportion

14 Hor explanation, see text.

130 ANNALS NEW YORK ACADEMY OF SCIENCES

of its species from the mainland by way of the other islands, it has the

smallest percentage of species which are not in the other Antilles. Porto

Rico, Hispaniola and Jamaica contrast with the Lesser Antilles (St.

Vincent) and Cuba in having a greater percentage of species which have

criginated in the Antilles and been passed to each other, but not to the

mainland.

Table X gives the percentages of the species occurring in a given island

and also on the mainland which are found in the several mainland divis-

ions. It is an attempt to discover the way species have moved in rela-

tively recent times. It probably concerns relatively recent movements

only, for the species have not changed enough to lead taxonomists to

give the mainland and insular lots different names. ‘T’o be sure, we

know that specific identity may be preserved for long geologic time and

TABLE X ¥

IL@Seeie |) 1B, Les Hisp | Cuba Jam

i Ul arene ne ee SS SP i |

Su AL omy. gg es Pee dee 3160 11.26 10.8 3.8. |) f4ss

Ch Aa SOM Sao ahs eee aan 5.2 (ea0) 2.7 6.3 fo

Ue Si Ory hee erie 6.9 e.) 10.8 30.0 0.0

Sort AL EG 3 As TIE eee aes 2 14.0 8.1 6.3 how

Cis, SANT BS ards tk OIE a ie 8.6 20.9 29.7 PA es 14.3

Saas WesS ec cee wee tte Bree 720 ie: ee 5 0.0

SP A CT IS INES ee ocn oie 2 Or gu acre 32.4 30.0 aT 2

Tore: eee 79 30 |. Goa 56.8 42.5 78.6

taal "Ge AN ite. ge ete DSO ie aye ee (220 a) Bears Sal7

Matyi; Auli keer 46.5 67 .4 18230) 1 Sas TAA.

experimental work has indicated that a species may arise not only sud-

denly but in several quite independent centers. However, most of the

species considered here probably arose in relatively recent geologic time

and each in a single center. Jamaica is troublesome because 33 (all but

14) of the species known from there do not occur on the mainland.

Cuba has 51 such species; Hispaniola, 39; Porto Rico, 26, and the Lesser

Antilles 110.

Except for the strictly South American species in St. Vincent and the

strictly northern species in Cuba a large part of the species which are

found in the various Antilles and also on the mainland are wide ranging

on the mainland, being found in all three of its divisions. This may be

due to the greater chance they have of getting to the Antilles (supposing

they originated on the mainland) because their mainland distribution

15 Hor explanation, see text. Based upon 58 species in the Lesser Antilles, 43 in Porto

Rico, 37 in Hispaniola, 80 in Cuba, and 14 in Jamaica.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 131

puts them on three sides of the Antilles or, as 1s more probable, they are

species which can live in a wide range of environments, as is indicated

by their wide geographic range, and so have found it easy to establish

themselves in the Antilles.

The Central American affinity is really slight. Although in the totals

it seems to be important, an examination of the more detailed part of the

table shows that this is caused by the groups in which it is joined with

either the northern or the southern division or with both. For example,

the 17.2 per cent. of the mainland species in the Lesser Antilles which

are credited to South America and Central America should probably be

eredited to South America as is indicated by the fact that 31.0 per cent.

are known on the mainland only from South America. The 5.2 per cent.

which are known from the mainland only in Central America are three

species which may have come directly from Central America but they

more probably occur in South America or did recently occur there.

Jamaica is the only island with a greater percentage of species which,

as far as we know, are strictly Central American, than it has of species

which seem to be confined to one of the other mainland divisions, but

this percentage is misleading also as it represents but one species.

Presumably these species have reached their island homes from the

mainland or have originated in the islands and spread to the mainland

since the various islands have been separated, if, indeed, the islands were

ever joined. The fauna of the islands at the ends of the Antillean chain

have their strongest affinities with that part of the mainland which is

nearest to them partly because of their geographical contiguity but partly

also because of resemblance of habitat, witness the Floridian affinity of

the fauna (and it is true also of the flora) of the sandy plains near Pinar

del Rio, Cuba, as compared with the more tropical character of the moun-

tains to the north of the city. If these species have spread themselves

in this way without the aid of actual land connection, is such connection

indicated in the older units, the genera ?

Jt is doubtless clear that if we take all genera and treat them as we

have just treated the species the result would be influenced by the in-

clusion of recently introduced species. Probably—but certainly not nec-

essarily—those genera which have species peculiar to the Antilles have

been on the Antilles longer than those whose only Antillean species occur

also on the mainland. Table XI gives the percentages of such presum-

ably old genera occurring in a given island and also on the mainland

which are found in the several mainland divisions. It is an attempt to

discover the way the fauna moved in relatively remote times.

132 ANNALS NEW YORK. ACADEMY OF SCIENCES

SABLE: DX

Lesser 12 deve Hisp. Cuba Jam.

S.A. only ee oe eee oil 6.9 9.4 10.2 5.9

Cy Ag tomy econ eee 2..8 3.4 6.3 4.1 Judi

U.S: sonlyes Se ee eee 4.2 3.4 0.0 Oak 0.0

S.. A. GC. DAC ee ee eons 21.1 ies DAR 18.4 11.8

CAL CUS ene errs 4.2 3.4 3.1 4.1 eT

Sa Ass SUNS hose ee ee ee ae 7.0 3.4 6.3 6.1 0.0

S. Ac Vel NeW Son een 39.4 62.1 53.1 51.0 53.0

otal) SE VAC S a ates 88.8 89.7 90.6 85.7 70.6

etal Gi Asks nc aeons 67 .6 86.2 84.4 tage 94.1

TotaleoUilesrs ss0 eee ae 54.9 72.4 62.5 3 (0.6

As with species, a large part of the genera which are found on the

various Antilles and also on the mainland are wide ranging on the main-

land, being found in all three of its divisions. A part of this is, of

course, due to those genera containing wide ranging species which also

have species peculiar to the Antilles. We also note that, as with species,

the strongest generic affinity of St. Vincent is with South America and

probably most of the 15 genera (21.1 per cent.) found in both South and

Central America got to or from Central America by way of South

America.

It has already been pointed out that the Porto Rican spider fauna is

largely recent. The data for genera as well as that for species indicates

that its affinity is probably strongest with South America. It would not

be profitable to discuss Hispaniola at length until we know more of the

fauna in the interior, but it is interesting to note that no strictly United

States genera have been found, and it is safe to predict that further study

of Hispaniola will show it to be even more South American than is Porto

Rico.

Cuba has a surprisingly small percentage of United States genera if

we leave out of account the wide ranging ones. The facts that 30.0 per

cent. of the species which are found on the mainland are not found on

the mainland south of the United States as compared with 6.1 per cent.

of presumably old genera, and that the figures for those not found north

of South America are 3.8 per cent. and 10.2 per cent. respectively indi-

cate that the older fauna has a stronger South American affinity than the

more recent. The meaning of this will be discussed presently.

There are only 17 genera in Jamaica upon which to base the percentages

given in Table XI. A strong affinity with Central America in ancient

16 For explanation, see text. Based upon’71 genera in the Lesser Antilles, 29 in Porto

Rico, 32 in Hispaniola, 49 in Cuba, and 17 in Jamaica.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 133

times is indicated, but as it is based on but 2 genera found on the main-

land only in Central America and Mexico, 2 found in these regions and

South America and 3 found in these regions and United States, it would

not be safe to place muth reliance on these data. Furthermore such

affinity does not demand a land bridge; it may merely be a result of geo-

eraphic contiguity and environmental similarity. If Jamaica has always

been isolated as it is now, or at least if its last complete submergence was

a long time ago and it has since been as isolated as it is now, we can

understand the apparent poverty of its fauna, the peculiarity of it and

its only shghtly stronger affinity with Central America than with the

other mainland divisions.

The impression all these things make on me is that the various An-

tilles may always have been as distinct as they are now; that they re-

ceived their spider fauna by slow “accidental” means, and that Porto

Rico has only recently been populated. How, then, are we to explain the

ancient character of a large part of its fauna and the curious relation-

ships with distant Old World localities? Also, why is the ancient portion

of the Cuban fauna more South American than the recent?

ORIGIN OF THE ANTILLEAN FAUNA

Before considering further the origin of West Indian spiders it would

be well to note their means of dispersal. Young spiders of nearly, if not

quite, all families are more or less given to “ballooning.” They will face

against the wind, and, elevating their spinnerets, spin a quantity of fine

silk usually in the form of a number of threads. These threads float in

the breeze and finally they are numerous enough or long enough to carry

their makers on an aerial journey. The length of such journeys would

seem to depend very largely on the strength and character of the wind.

McCook has attempted to show by its distribution that Heteropoda vena-

toria has circumnavigated the globe. It is true that the range of this

species corresponds “with remarkable exactitude” to the belt over which

the Trades blow, but we may accept his proof that the species has not

been distributed by commerce without adopting his suggestion that its

cosmotropical distribution is due to its ballooning habit. He cites Dar-

win’s note about the “Beagle” being boarded by “flying” spiders when

sixty miles from land and adds a report which is similar except that the

latter ship was more than two hundred miles from land. It must be ad-

mitted that wind may be a very efficient factor in the distribution of

many organisms, and it should be noted that when spiders go on such

journeys they often go in. swarms, so that it would not be unlikely that

opposite sexes would land near enough each other to continue the species.

134 ANNALS NEW YORK ACADEMY OF SCIENCES

However, there is more to the distribution of spiders than wind. We

need an explanation of the remarkable discontinuities which have been

pointed out.

Ocean drift is frequently brought in to explaim distribution and it has

probably been the effective agent in many instances. The opponents of

such an idea forget the long ages in which accidental drift has had a

chance to work. However, the ease with which large numbers of spiders

take to the air every year makes recourse to the small numbers that may

make successful voyages on driftwood unnecessary.

Then there is man. Hardly a ship sails from port without araneid

stowaways, and inland shipments of freight also carry their quota, but

these are nearly always a certain few species of which only those that live

about man’s dwellings are likely to become established. None of the

“tarantulas” which come in nearly every shipment of bananas have be-

come a part of the New York fauna. Furthermore it is very unlikely

that the distribution of most spiders has changed much since man began

to sail the seas. The study of their movements must go far back of that.

One takes up the question of land bridges with something akin to a

groan since opinion on the matter is so diverse and the evidence, whether

pro or con, about a given bridge is often so slight. Paleontology offers

little direct evidence as to the ancient movements of spiders since so few

fossil spiders are known. Several have been described from the Carbon-

iferous of both hemispheres and probably at least one genus (Arthroly-

cosa) was even then found in both Europe and North America. The

only living genus of the type of spiders which was apparently common

in Carboniferous is Liphistus. It is found now only in the islands of

Pinang and Sumatra.

Spiders with unsegmented abdomens, that is all living spiders except

Lipistius, may not have arisen until the Mesozoic, but, if so, evolution

was fairly rapid, for most of the Oligocene and Miocene spiders belong

to present-day families and even genera.

R. T. Pocock** gives an interesting analysis of the ancient movements

of spiders. A few of the present-day distributions given by him differ

from those given by other authorities, but none of the differences which I

have noticed would materially change his argument. In his section on

the “Distribution of some of the Families of Arachnomorphe that were

represented in the Oligocene Period” he mentions seventeen genera found

in amber which are still hying. While it is perfectly true that “although

since the Oligocene these Spiders have had the same time for dispersal,

they nevertheless differ greatly in their distribution,” the implication that

Proc. Zool. Soc. London, 1903, I. pp. 349-568.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 135

their present distribution is a measure of their success in extending their

range does not necessarily follow. It is altogether probable that spiders

fairly covered the earth in Oligocene times, and the few that were im-

prisoned in Baltic amber tell us nothing more than that they lived in that

region, among others, at that time. It is pleasing to note that Pocock

did not think it necessary to throw a bridge from continent to continent

on the shortest line (according to Mercator’s projection) between any two

portions of a present-day discontinuous range. He naturally found con-

siderable evidence of transfer in Arctic regions** and he also used, rather

freely, Antarctic connections between South America, Africa and Aus-

tralia.

F. Dahl*® has considered the distribution of spiders and concluded that

the Antarctic connection is improbable. Not all of his arguments seem

to be well founded. For example, one can not consistently uphold the

“Telict” idea and then combat the Antarctic connection on the ground

that so few relicts are found on the Antarctic islands. Most of the spi-

ders, for the explanation of whose distribution an Antarctic connection

might be desired, are tropical. It is therefore not surprising that re-

licts are not found on Antarctic islands.

Ité is no more than ordinary common sense to favor the simplest ade-

quate explanation of a problem. It is unnecessary to review the volu-

minous literature concerning Antarctic connections between the three

southern continents. It is admitted that many facts favor such connec-

tions, especially the one between South America and Australia, and that

nothing has been, or is likely to be, brought out which will absolutely

disprove it. However, if a simpler explanation than the general eleva-

tion of more than 3,000 meters required to connect South America and

Australia, but which leaves Africa still to be accounted for, is adequate

it would seem to be preferable. The final court of appeals is, of course,

fossils, especially those in Antarctica, but even if numerous fossils are

found in Antarctica and they are seen to be similar to those found in the

southern portions of the other continents it will not prove actual con-

nections. It will merely show that there has been an interchange of

fauna—a thing not at all unknown between absolutely unconnected land

areas and a thing for which there was a vastly longer time than we

ordinarily have in mind since it requires a strong effort for us to bring

ourselves to thinking in terms of millions of years.

18 The theory that practically all Tertiary migration was by way of the Arctic regions

has been set forth most clearly by W. D. Matthew. It is amplified and convincingly dis-

cussed in his recent paper on “Climate and Evolution,’ Annals N. Y, Acad. Sci., Vol.

AXIV, pp. 171-318. 1915.

*% Zool. Anzeiger, vol. XXXVII, pp. 270-282. 1911.

136 ANNALS NEW YORK ACADEMY OF SCIENCES

In addition to, and partly because of, our relative ignorance of ancient

spiders, we known but little of araneid phylogeny. We do know that

spiders were already fairly well distributed, at least in the northern

emisphere, in the Carboniferous. Modern families and even genera

were well differentiated in Oligocene. Therefore, as far as distribution

problems are concerned, the present-day distribution ‘of primitive genera

in a given family or even primitive species in a given genus is of as

much, or more, importance as the present-day distribution of primitive

iamilies. We unfortunately lack a sufficient knowledge of the compara-

PNAS aicrt

ii Re Sica maw iS * 7

RE TNE

= es Bey ees OS

ll R

ee CAL Me]

Fic. 5.—Distribution ef Archeide

A, Fossil (amber) Archea; B, Living Archea; C, Mecysmauchenius.

tive anatomy of spiders to enable us to use such data in the solution of

the problems of distribution. In fact, widely discontinuous distribution

is, at present, the best indication we have of relative antiquity and its

use is rendered hazardous by reason of the possibility of polyphyletie

origins—a possibility which has been rendered more probable by recent

work in experimental evolution.

An analysis of the distribution of the world’s spider fauna would be

out of place in this paper and only a few points which seem to have a

bearing on the West Indian problem will be taken up.

The family Archeide is an interesting one in this connection (see

figure 5) since Baltic amber contains a genus (Archea) of which the

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 13

only known living representatives are in Madagascar. Furthermore, the

only other genus of the family is confined to Patagonia. Had it not

been for the fortunate find in Baltic amber we would not know that the

family ever occurred in the northern hemisphere and a land bridge from

Patagonia to Madagascar in the most direct way compatible with sea-

bottom contours would receive additional strong support. As it is, it

seems more probable that this was once a fairly widespread family and

it certainly occurred in northern Europe. We know that prior to the

glaciers the climate of the North Polar regions was mild or even tropical

Fic. 6.—Distribution of certain genera known from Baltic amber

1. Segestria; 2, Dysdera; 3, Eresus; 4, Amaurobius; 5, Archaea; 6, Agelena; 7,

Anyphena. Only certain points in some of the ranges are indicated. The lines

represent possible routes of dispersal.

and hundreds of circumpolar species point to the interchange of fauna

between Eurasia and North America, where land bridges, if they ever

existed, would not need to be long, so that it really seems quite probable

that the genera of Archzide which are found in Madagascar and Pata-

gonia respectively are merely remnants either of a formerly widespread

family or of a northern family which was driven south by competition

with new forms. Such a movement is supported by mammalian fossils

and seems more likely than a South Polar connection.

Reference has been made to Pocock’s list of seventeen recent genera

found in Baltic amber. Of these we cannot consider here Aranea and

138 ANNALS NEW YORK ACADEMY OF SCIENCES

Zilla because their taxonomy is too unsatisfactory. Drassus has been

split up by authors but even one of its parts is practically cosmopolitan.

Tetragnatha, Tegenaria and Philodromus are cosmopolitan, even includ-

ing frigid regions. Olubiona is found throughout most of the temperate

and tropical regions ; Nephila and Sprassus are cosmotropical; and Thom-

isus is found throughout most of the continental, at least, Old World.

Certain points in the ranges of the other seven genera are shown in figure

G, only those most distant from the Baltic together with several inter-

mediate stations being indicated. The supposition that all these seven

Pcs Paes

eRe Sei sea

LCC eR

Hint asc

ot eee

: ee

5 z

2) |

{

e |

G 1 ES

Hic. 7.—Distribution of Segestriine

Lines refer to Segestria and dots to Ariadna.

cenera originated in the Baltic region in, say, the Eocene would be ab-

surd, but supposing that they did, would not the several million years

since then, including as they do a long space of mild or tropical condi-

tions in the Arctic, have been sufficient for the dispersal of these genera

more or less along the lines indicated in the figure? I feel that even

the rather well authenticated land connections in the North Polar region

would not be necessary, for we must remember that the distances are

much shorter in the polar regions than they appear to be on a Mercator’s

projection. Certainly, there seems to be no need for Antarctic connec-

tions. The distribution of Archea has already been mentioned. The

only other genus of special note is Segestria. The discontinuity of its

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 139

distribution is toned down by a consideration of Ariadna, a genus which

with it forms the subfamily Segestriine. Thus Segestria in New Zealand

is linked with Asia by Ariadna in Australia, Sumatra and Ceylon. See

figure 7

By selecting certain genera or groups of genera of spiders it would be

easy to make quite a list in favor of a land bridge from South America

to Africa and elsewhere such as that shown in figure 8 which is after

A. E. Ortmann’s idea of certain of the connections in Upper Cretaceous

times. For example the following genera might be cited: Cyatholipus

<i S| 3 malo eed E ce ?

New ¥ ; ~~ E oF

ARMac ie

Sf aes

iaatedantgaataaat

ere

Fie. 8.—One of the hypothetical Upper Cretaccous land masses

and Drymusa, Antilles and South Africa; /schnothyreus, Antilles, West

Africa, Ceylon and Philippines; Caloctenus, Antilles, West Africa and

Malasia; Dyschiriognatha, Antilles, Venezuela, Egypt, Ceylon, Borneo

and Japan; Opopea,?® Antilles, Peru, Colombia, Venezuela, Africa,

Arabia, Ceylon and Philippines; Anapis, Antilles, Venezuela, Brazil,

northwest Africa and New Caledonia; Ogulnius, Antilles, Brazil and

Ceylon ; Hpisinopsis, Antilles, Peru and Malay Peninsula; Theotima and

Accola, Antilles, Venezuela and Philippines; Beata, Antilles, Mexico to

Brazil and Africa; Syrisca, Colorado, Utah, Texas, Brazil and Africa;

Oxyopeidon, Arizona to Panama, Antilles, East Africa, India and Indo-

7? Indeed the range of one species (O. deserticola Simon) is St. Vincent, Venezuela,

Egypt, Arabia and Philippines.

140 ANNALS NEW YORK ACADEMY OF SCIENCES

China; Ochyrocera, Pacitic coast states, Mexico, Antilles, Venezuela,

Brazil, Ceylon and tropical Asia; and Physocyclus. California, Arizona,

and New Mexico to Colombia, Guiana, Antilles, Africa and tropical Asia.

All of these are on this particular hypothetical land mass and not else-

where, as far as is known, except possibly several Peruvian or Brazilian

localities. They were picked in a rather random fashion and only Antil-

lean genera were included. ‘Taking the world as a whole the lst could

be increased beyond the patience of the reader, but what would it prove?

In general, it proves that it is possible to select a long list of genera and

even some species which accord with a certain hypothetical land mass.

In particular, the tropical parts of the area selected here includes some

of the most important preserves of ancient types. We may leave poly-

phyletic origin of genera out of account not only because its occurrence —

is unproven but because it would not influence the case since a similar

base from which to get the separate origins would be required and hence

the problem would be merely restated, not changed. We have then to

decide as to which is more reasonable: (a) forms migrated over this

hypothetical Jand mass, which crosses the Indian Ocean and the Atlantic

at the Equator, and then died out or have not been found except in cer-

tain favored spots on this mass; or (6) forms spread over land which

now exists and, if you please, the shorter land connections but not the

oceanic ones, and have died out or have not been found except in certain

favored spots. A very readable and fair summary of the arguments in

favor of the first alternative is given in Scharfi’s “Distribution and

Origin of Life in America.” He also favors a second Atlantic land mass

running from the Antilles to the Mediterranean region with an offshoot

to Bermuda and Nova Scotia. A list of spiders may be compiled from

the taxonomic part of this or other papers which will accord with this

bridge. The bridge was supposed to have existed in early Tertiary and

to have also connected northeastern America by way of the Great Lake

and Rocky Mountain regions with the southern tip of South America.

Most of South America was under water; eastern Brazil and part of the

present Atlantic bed formed a large island, while part of Ecuador and

Peru formed the tip of a peninsula from the main Jand mass which lay

in what is now the Pacific and included the Galapagos, a strip running

across the middle of Central America to the Antilles, and in its north-

western sweep reached from California to Hawaii, but just before it got

to western Canada it turned west, missed Alaska and connected up with

Asia. The thing is rather complicated but certain spiders might have

traveled the route. Note in this connection that several genera and

groups of genera are known from northeastern North America and Pata-

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS 141

gonia but are only poorly or not at all represented elsewhere. However,

is it necessary? I think not. Even with ocean bottoms raised to form

connections where connections are wanted and present continents sunk

to explain the absence of forms where they are not found, one must fur-

ther make all the assumptions required for the theory that distribution

has been accomplished by present-day land masses. Forms either origi-

nated in America and passed to the Old World by way of the Arctic or

the movement was in the reverse direction or, as is more probable, both

are true. If geology demonstrates that there were land connections

there, the passage will be seen to have been easy. However, there is no

necessity for such connections since the distances are short and the time

was long. The more recent American forms probably originated, for

the most part, in America. The cutting off of the Arctic route by the

lowering of the temperature has temporarily, at least, prevented natural

interchange between the two hemispheres. Those recent forms which

originated in the north are adapted to temperate conditions (which in-

cludes more than mere ability to withstand cool winters) and they have

replaced in the north the old fauna which was adapted to more tropical

conditions. In the south there are numerous remnants of this old fauna

together with recent offshoots of it. The only assumptions necessary for

this theory are really statements of fact, namely, that the climate of the

Arctic was at least mild in the Tertiary and that many formerly wide

ranging forms are now restricted in their distribution.

SUMMARY

To summarize what seems to be the facts concerning West Indian spi-

ders: there has been considerable movement between the individual

islands and also between the mainland and the islands, especially at the

two ends of the island chain, even in recent times when the islands were

separate from each other and from the mainland. It is therefore unnec-

essary to suppose that such connections ever existed. Ancient forms

- have had a longer time to reach the islands than the more recent ones,

they were adapted to a tropical environment, and the insular character

of the area has protected them, hence a large part of the fauna consists

of relicts as is shown by the relationships with South Africa, Madagascar,

Ceylon, Australia and the Philippines. Recent forms are now mingling

with and replacing the older forms.

142 ANNALS NEW YORK ACADEMY OF SCIENCES

BIBLIOGRAPHY

(For references to special species see Petrunkeyvitch’s Catalogue)

BANKS, NATHAN: Some Spiders and other Arachnids from Porto Rico. Proce.

U. S. Nat. Mus., XXIV, 207-227. 1902.

: A List of Arachnida from Hayti, with Descriptions of New Species.

Proc. Acad. Nat. Sci. Philadelphia, LV, 340-345. 1903.

: Arachnida of Cuba. Secretaria de Agricultura, Comercio y Trabajo

de la Republica de Cuba. Estacion Central Agronomico, 2nd Report, Part

II (English Edition), 150-174. 1909. .

Lutz, FRANK E.: The Distribution of Occidental Spiders. Science, N. S.,

XXXVII, 567-568. 1913.

PETRUNKEVITCH, A.: A Synonymic Index-Catalogue of Spiders of North, Cen-

tral and South America, with all Adjacent Islands, Greenland, Bermuda,

West Indies. Terra del Fuego. Galapagos, ete. Bull. Amer. Mus.- Nat.

Hist., XXIX, (91 pp. 1981:

Srtmon, EuGENE: Histoire naturelle des Araignées. 2 Volumes. Paris, 1892-

1903.

ScHwartz, E. A.: Proc. Ent. Soc. Washington, VI, 147-148. 1904.

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS

Acacesia, 91, 115

Accola, 77, 111, 139

Actinopodins, 75

adansoni, 105

Agelenide, 100

agilis, 76

Agobardus, 103, 110

aissus, 97

albopalpus, 103

Alcimosphenus, 88, 111, 115, 117, 120

americanus, T8, 95

Anapis, 111, 139

anastera, 91

anglicana, 87

anne, 107

enormalis, 103

antiguensis, 97

antillana, 88

antillanum, 84

antillanus, 106

Anyphena, 98, 99, 116

arabesca, 91, 92

Arachnomorphe, 134

Ar@ned, 137

Araneus, 87, 91, 110, 127

Archea, 108, 136, 137, 138

Archeidz, 108, 136, 137

Arctosa, 102

arenatus, 91

argentata, 90

Argiope, 90

Argiopid, 78, 86, 87

Argiopineg, 90, 95

argyra, 88

Argyrodes, 84

Argyroepeira, 88, 89

' Ariadna, 81, 138, 139

armata, 94

aurmatus, 104

Artema, 83, 111, 113, 121

Arthrolycosa, 134

asperatus, 95

athletica, 75

atlanta, 83

atlantica, 102

Atypidze, 108

INDEX

audax, 104

aurantia, 90

aurantius, 104

aurata, 104

aussereri, 102

Avicularia, 76

Avicularids, 75

Aviculariins, 76

Aysha, 98, 114

badia, 102

bajula, 80

balaustinus, 92

banksi, 107

Barychelinsz, 76

Bathyphantes, 87, 117

Beata, 111, 139

bellicosus, 105

bellulus, 95

benjaminus, 92

bicolor, 81, 97, 100

bifurcatus, 88

bigibossa, 88, 89

bispinosius, 92

bivittatus, 106

Blechrosceleze, 83

Blechroscelis, 83, 110

Bolostromus, 111

brunneda, 87

Bythocrotese, 103

Bythocrotus, 103, 115, 120

cesia, TT

Callilepis, 82, 117

Caloctenus, 111, 139

cancellatus, 84

cancerides, 77

cancriformis, 94

canestrinii, 94

capitatus, 104

Caponiide, 81, 109

Caponina, 82

caroli, 90

Castaneira, 99

cavernicolus, 86

celer, 96, 97

143

144 ANNALS NEW YORK ACADEMY OF SCIENCES

cephalotes, 103

Ceratinella, ST. 117, 121

Ceratinopsis, S7, 117

Chiracanthium, 98

cincta, 82

cinerea, 102

clavipes. S9

Clubiona, 98. 138

Clubionidz, S82, 97

Clubioninz, 98

Diphya,. 89

Diplura. 77

Diplurinz, 77

directa, 91

Dolichognatha. SS

Dolomedes, 101. 116, 120

domestica, 100

Drassidz, $2

Drassus. 138

Drezelia, 91. 117

coccinea, S82, ST , Drymusa, 80. 111, 113. 120, 139

cockerelli, 106 Dysderidez, $1, 108

Compsodecta, 103 Dysdertma, 81. 111. 113, 121

conchiea, 91 Dyschiriognatha, 111, 139

concolor. S9

conjormis, T5

Corinna, 100, 116

Corinninz. 100

Corinnomma, 110

coronotus, 107

Corythalia, 104

crassicauda. 90

crewti, 92

Crotolaria, 98

Ctenex, 99

Cteninz, 99

Ctenizinz, 75

Ctenus. 99

cube, 99

cubana, 76, 92, 94. 97. 98

cubanus, 76, 79

Cupiennius, 99, 117

cursor, 76

Cyatholipus. 88, 119, 120, 139

Cyclosa, 90

Cyrtopholis. 76

Cyurtophora,. 90, 115

defioccatus, 107

Dendryphantes. 104. 110, 127

dentipes, SS

derhami, 100

descripta, 100

deserticola, 159

Diewa, 96

Dictyna, 79, 117

Dictynidz, 79, 10S. 110

digitata. 77

Dinopinz. 7S

Dinopis. 78

dubius, 101

echinatus, 96. 116, 120

Edricus, 90, 113

Blliea, 82, 116, 117

elegantissima, 104

elongata, 8S

élongatus. S

Epecthina, 95

Epecthinula. 95. 119. 120

Epeira, 91, 92. 98

Episinopsis. 111, 139

Eresidz, 108

érichsonit, 77

Friophora, 92, 94

EBrissus. 96

ernsti. 100

EBugnatha, SS. 114

Eurupelma, 75. 76. 117

EBustala, 91

Eustiromastiz. 105

Eutichurus. 98.115, 117

Evophrydee. 105

Erophrus. 105

jastuosa,

jeroz, 98

jictilium, S4

Filistata, 80. 119

Filistatide. SO

fiavidus, $4

fiaromaculata. 94

foliata, 91

forcipata. 94

formica. 107

frondeum, S+

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS

Fuentes, 106

juesslyi, 84

fusca, 80, 102

fusco-vittata, 91

Gasteracantha, 94

gastercanthoides, 95

Gea, 90, 117

geniculatus, 78°

Geodephaga, 60

geometricus, 85

glaucus, 83

globosa, 93

globosus, 83

Glyptocranium, 94

Gonatium, 86

Gongylidiellum, 86

gracilipes, 98, 100

gracilis, 88

granadensis, 92

grisea, 82, 102, 103

grossa, 85

guanabacoe, 82

gundlachi, 92

guttatus, 85

Hahnia, 100, 111, 115, 121

haitiensis, 99

Hamatativa, 102

Hapalopinus, 76, 116. 117, 120

Hasarius, 105, 110

haytiensis, 105

Hedana, 96

Hedypsilus, 83

heptagon, 90

Hersiliidz, 108

Heteropoda, 97, 133

hexacantha, 94

hibernalis, 80, 99

hilaris, 94

hirsutus, Té

horrida, 94

hortorum, 89

humilis, 100

Hyctia, 105, 117, 118, 121

Hypochilidx, 77, 108

Hyptiotes, Ts

foves, 105, 115, 116

illustris, 107

imnvaculata, 99

incerta, 106

incertus, 92

inclusum, 98

inocuus, T6

insignis, SO

insulanus, 98

insularia, 100

insularis, 82, 97, 99, 102

-interfector, 95

interruptum, 85

inutilis, 107

Tsaloides, 96

Ischnocolus, 76, 110

Ischnothele, 77, 119, 121

Ischnothyreus, 110, 139

jamaicola, 76

Janulus, 111

keyserlingi, 82, 99

laboriosa, 88

labyrintheus, 92

lata, T7

laimnia, T8

Larinia, 91

larvatus, 84

Lasiodora, T6

Latrodectus, 85

Leptonetidz. 81, 108

Leucauge, 88

licinus, 88

lineatipes, 80

Linyphia, 87, 115

Linyphiidz, 86

Linyphiinex, 87

Liocranine, 99

Liphistiidz, 75

Liphistius, T5

Liphistus, 134

Lipistius, 134

Lithyphantes, 86, 117

locuples, 104

longipes, SO

Lycosa, 102

Lycosidx, 101

Lyssomanee, 105

Lyssomanes, 105, 106

146

macrura, TT

mactans, 85

maculata, T5

maculatus, 97

Meevivese, 106

major, 104

malvernensis, 99

Mangora, 90, 117

marginella, 101

maritima, 98

Marpissa, 106

Maraxia, 93

Mecolesthus, 83, 113

melanocephalus, 106

melanognatha, 106

mendicus, 104.

Menemerus, 106

Mesothele, 75

Meta, 88, 89

Metacyrba, 106, 115

metallica, 104

Metaphidippus, 104, 105

Metazygia, 94

Metepeira, 92

mexicanus, 92

Miagrammopes,-79, 111, 113, 115, 116;

ele. ial

Miagrammopins, 79

Micariine, 99

Micrathena, 94

Microctenus, 99

Microneta, 87

Miginze, 75

Mimetide, 95, 108, 110

Mimetus, 95

miniatus, 105

minutissima, 95

Misumena, 95

Misumenins, 95

Misumenops, 95, 96, 116, 120

Misumessus, 96, 116, 120

Modisimus, 83

morgani, 107

Mygale, 75, 110

Myrmarachne, 106, 117

Myrmecium, 106

Mysmena, 86, 117, 121

nauticus, 92

navus, T9

ANNALS NEW YORK ACADEMY OF SCIENCES

Néoscona, 91, 92, 93

néotheis, 93

Nephila, 89, 138

nephile, 84

Nephilinz, 89

nidulans, Td

Nilacantha, 106, 115, 120

nitidum, T6

nomina, 83

Nops, 82

nubila, 80

nuda, 83

oaxacensis, 93

oblongus, 96, 97

obtuspina, 94

Ochyrocera, 81, 140

octavus, 104

octopunctatus, 105

oculata, 90

Gicobtide, 79, 108, 110

Gicobius, 79, 113

Ogulnius, 111, 139

Olios, 97

Onoculus, 96

Oonopide, 81, 108

Oonops, 111

Opoped, 81, 111, 131; 1210 igs

opulenta, 85

Otiothops, 82

Oxyopeidon, 103, 111, 113) tite

Oxyopide, 102

Oxyopes, 103

Oxysoma, 98, 116

Pachylomerus, 75, 119

pallens, 98

pallescens, 88

pallida, 94

pallidus, 103

Palpimanide, 82,

Palpimanine, 82

paradoxa, 84

parallelus, 106

Paraphidippus, 104

Paratropidine, 75

Pardosa, 102, 113

parietalis, 79

Parnenus, 104

parvula, 99, 107

109

LUTZ, LIST OF GREATER ANTILLEAN SPIDERS

paykulli, 107

peckhami, 108

Peckhamia, 106, 117

pegnia, 93

Pellenes, 106, 118, 119, 121

Pensacolez, 103

pentagona, 96

perplexa, 107

perplexides, 107

perplerus, 93

perpusilla, 98

Peucetia, 103

philangioides, 83

Phidippus, 104, 105

Philodromine, 96

Philodromus, 138

Pholcidz, 82

Pholcinez, 82

Pholeus, 83. 110

Phormictopus, TT, 114

Physocyclus, 83, 140

picata, 106

pictipes, 106

pikei, 105

Pisauridex, 101, 110

piscatoria, 88

placida, 91

Platoridx, 108

Plexippus. 107

plumipes, 78

poeyi, 103

portoricensis, 102

pratensis, 95

pretiosa, 99

Prodidomidz, 108

prompta, 91

Prostheclina, 107, 113. 114. 115, 116,

mer Ais, 179, 120

proximus, 105

prudens, 105

Psechridze, 108

Pseudosparianthis, 97

punctulata, 102

pygmea, 104

quadripunctata, 85

rand, 103

ravida, 98

regius, 105

regnyt, 89

republicanus, TS

retiarius, 107

retusca, 98

Rhomphea, 84

rufipes, 84, 85

rufopunctata, 94

rugosus, 96

Saitis, 107, 113, 118, 119, 120. 121

Salticidz, 103

Salticus, 103

Schizopelma, T7

scoparius, T9

Scopelobates, 76, 115, 120

Scytodes, 80

Scytodidze, 80

Segestria, 137, 138, 139

Segestriinz, 138, 139

Selenopins, 97

Selenops, 97

sellata, 90, 104

semicincta, 87

Senoculidz, 108

seperatus, 105

septenmaculatus, 86

Sergiolus, 82

sericatus, 93

sericeus, 76

serripes, 83

sexserrata, 94

Sicariidz, 80, 108

Singa, 92

signata, 107

signatus, 83

Simonella, 107

sloanci, 94

Smeringopus, 83, 117

Solenodon, 125

solitaria, 81

Sparassine, 97

spherula, 85

spinicrus, T6

spinosa, 78, 96

Spintharus, 84

Sprassus, 138

stellatus, 93

Stephanopsins, 96

Stephanopsis, 96, 111, 117, 121

Stichoplastus, 76, 113

Stoidis, 104, 114

147

148 ANNALS NEW YORK ACADEMY OF SCIENCES

Stothis, 76 tredecium, 85

striata, 98 triangularis, 85

studiosum, 84 Trichopelma,. 76, 115

suavis, 108 trifasciata, 90

sylvana, 107 trituberculatus, 84

Synemosyna, 107 trivittata, 92

Synemosyneze, 107 tuberculata, 89

Syrisca, 82, 99, 116, 117, 121, 139

undecimtuberculatus, 94

tarantulas, 76, 134 undecimvariolata, 92

tauricornis, 93 Uliodon, 99

taylori, 105 Uloboridz. 78

Tegenaria, 100, 117, 138 : Uloborinz, 78

Temimius, 82, 99 Uloborus, 78, 79

tenuis, 98 Utetheisa, 98

tenuissima, 99 utriculata, 104

tepidariorum, 84 varia, 87

tetracantha, 94 variegatus, 82

Tetragnatha, 88, 138 variolatus, 94

Tetragnathine, 88 velox, 98

Teutana, 85, 110 venatoria, 97, 107, 133

Thanatidius, 101, 116, 117 venusta, 89

Thaumasia, 101, 116 vernalis, 108

theis, 938 Verrucosa, 91, 92

theisti, 93 viaria, 107

Thelechoris, 77 vicina, 88

Theone, 111 viridans, 96, 103

Theotima, 81, 111, 139 viridis, 106

Theraphose, 77 vituperabile, 84

Theridiidz, 84, 86 volatile, 85

Theridion, 84, 85 vulgaris, 93

Theridionexus, 86, 119, 120

Theridula, 85, 117 Wagneriana, 93, 94

Thomisidx, 95 Weata,: 108, 118, 415, 147

Thiodina, 107, 117 walckeneri, 82, 90

Thiodine, 106, 107 wilderi, 89

Thomisidz, 95 wistariana, 89

Thomisus, 138 wittfeldz, 94

Tibellus, 96, 117 Wulfila, 99, 114

Tillandsia, 104

tipuloides, 83 Zenodores, 104

Tobias, 96, 115 . Zilla, 138

toussaintii, 96 Zodariide, 108

Trachelas, 100, 116 Zoropside, 108

translatus, 107 Zygoballezx, 108

Trechona, T5 Zygoballus, 108

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- VOLCANIC DUST VEILS AND CLIMATIC

Se VARIATIONS

HENRYK ARCTOWSKI

NEW YORK » J ational

PUBLISHED BY THE ACADEMY

21 June, 1915

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[ANNALS N. Y. Acap. Scr., Vol. X XVI, pp. 149-174. 21 June, 1915]

VOLCANIC DUST VEILS AND CLIMATIC VARIATIONS!

By HENRYK ARCTOWSKI

(Presented in abstract before the Academy, 7 December, 1914)

CONTENTS

Page

NRE RTREME IS 0 ned Nn AT ee ati Soe ee alan, Sorta el ashes aha. sé 6S Bla a Cee 149

Volcanic eruptions of the years 1883, 1902 and 1912.................... 150

oie Pomel] ITE TOCOEUS so fc ous sic <tlniisea + sis dis a oe ce ws eed « 156

Temperatures at Port Darwin and some other stations.................. 158

memperacures at Wori-de-France, Martinique... ......-......- cc ccscccacces 162

- Temperatures at Para, Cayenne and the West Indies.................... 164

ere OUSCEVCE 1° ISK ts hee oe se de See Cee eee tee ee. 166

Temperature variations at Stockholm and Batavia....................-.- 168

Sellcuees and atmospheric temperature. ..:.. <<<... cece se oeceeecnees 169

a UI EDI Ie os oe fa ta cist he acta ain 2s ie Ride mite: Wf Sieg, © aiaiteje u,2 's8,8, 8.9 6 Gumnes 174

INTRODUCTION

The series of overlapping yearly means of temperature, expressed

graphically, show most characteristic crests and depressions. In the case

of tropical stations, in particular, the crests of the curves are very regular

and occur at intervals of two to three years,” practically at the same time

all around the world.

As a general result of a detailed study of the temperature data for the

years 1900-1909, for Europe, Greenland and North America, I have

found some striking correlations between these equatorial variations and

the more complicated variations of temperate and arctic regions.*

In another study of all available temperature data for the years 1891-

1900, I have shown that terrestrial atmosphere at the earth’s surface

was warmer in 1900 than in 1893 by at least 0.6 °C.* On the maps

representing the geographical distribution of the departures of annual

means from the normals or from the quasi-normal values of ten-yearly

means, the areas of positive departures have been called thermopleions

1 Manuscript received by the Editor, 4 January, 1915.

*HeNRyYK ArCTOWSKI: “The Solar Constant and the Variations of Atmospheric Tem-

perature at Arequipa and some other stations,’’ Bull. Am. Geog. Soc., vol. 44, p.598. 1912.

3 HENRYK ARCTOWSKI: “A Study of the Changes in the Distribution of Temperature

in Europe and North America During the Years 1900 to 1909,’ Annals N. Y. Acad. Sci.,

vol. 24, p.39. 1914.

_ * HENRYK ARCTOWSKI: L’enchainement des variations climatiques. Bruxelles, 1909.

(149)

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150 ANNALS NEW YORK ACADEMY OF SCIENCES

and the areas covered by negative departures antipleions. On the curves

of overlapping means, the crests correspond to pleions and the depressions

correspond to antipleions. I have presumed that the excess of pleions

over antipleions, corresponding to pleionian crests of equatorial stations,

may be due to an increase of the solar constant.°

Recently, many papers have been published about the influence of

voleanic dust on meteorological phenomena, on atmospheric temperature

in particular. I will simply cite the exiensive researches of W. J.

Humphreys,® C. G. Abbot and F. E. Fowle* and of H. H. Kimball®

Humphreys, in particular, does not hesitate to admit “that volcanic dust

inust have been a factor, possibly a very imporiant one, in the production

ef many past climatic changes.” |

The hypothesis ascribing the origin of climatic variations to the pres-

cnce of volcanic dust veils in the higher atmospheric layers is a very

plausible argument against my supposition that the changes in terrestrial

temperature are due to cosmical causes. Before going any farther in

my researches on the mode of formation and the dynamics of pleionian

variations, it was therefore necessary to find out to what extent one may

be justified in supposing that the antipleionian depressions of tempera-

ture are simply caused by the presence of volcanic dust veils. ;

In this paper I will show that though, in some cases, volcanic eruptions

may have influenced atmospheric temperature very greatly, this cause of

climatic variations is purely accidental and secondary, and that the

pleionian phenomenon is independent, in its cause, of the occurrence of

volcanic dust veils.

VoLcanic ERUPTIONS OF THE YEARS 1883, 1902 AnD 1912

Although we know, from geological records, that the volcanic activity

of the earth’s crust has undergone important changes, and although the

historical data indicate that seismic and volcanic phuiwivacied have varied

in intensity and frequency, our knowledge of these fluctuations, of long

as well as of short duration, is most unsatisfactory.

Papers on a possible relationship between sun-spots and volcanic phe-

5 HENEYK ARCTOWSKI: “About Climatic Variations,” Amer. jour. Sci., vol. 37, p. 3053..

1914.

*w. J. HumpuHeers: “Volcanic Dust and Other Factors in the Production of Climatic

Changes and Their Possible Relation to Ice Ages.” Bull. Mount Weather Observatory,

vol. 6, p.1. 1913.

™C. G. Apsot and F. E. Fowie: “Volcanoes and Climate.” Smiths. Misc. Coll., vol. 60,

No. 29. 1913.

§ Heepeet H. Kimpatt: “The Relation Between Solar Radiation Intensities and the

Temperature of the Air in the Northern Hemisphere in 1912-13." Bull. Mount Weather

Observatory, vol. 6, p. 205. 1914.

ARCTOWSKI, VOLCANIC DUST VEILS 151

nomena have been published by E. Kluge,® Joseph O’Reilly,*® Charles

Zenger,"' H. I. Jensen’* and others. As will be seen later, a very sug-

cestive conclusion may also be derived from the annual frequency list of

voleanic eruptions compiled by Leo Kelley. If, therefore, some striking

coincidences between changes of atmospheric temperature and the in-

crease of frequency or violence of volcanic eruptions exist, the existence

of these coincidences is a very inadequate argument in favor of the

hypothesis that the variations of temperature are due to volcanic dust

veils, since both phenomena, the volcanic paroxysms as well as the coin-

cident temperature changes, may be the effect of some common extra-

terrestrial factor.

In this study, however, only the volcanic eruptions of an explosive

character have to be taken into special consideration. Moreover, it is

only when volcanic dust has been projected in great quantity above the

ordinary elevation of the cirrus clouds, that is to say to an altitude of

8,000 m. or higher up, that this dust could remain in suspension long

enough to be spread out all around the globe by the winds of the lower

stratosphere.

In the case of the famous Krakatoa eruption of 1883, the optical

phenomena produced by the volcanic dust veil have been studied very

extensively. For reference it will be sufficient to cite the reports of

Rh. D. M. Verbeek** and of the Royal Society.**

Krakatoa is a small island between Sumatra and Java and lies 6° S.

lat. The explosion occurred on August 27, 1883. The main sky phe-

nomenon produced by the dust went around the world in fifteen days

irom east to west along the equator, spread out north and south, was

observed in the Gulf of Mexico by the end of September and all over the

States in November. The extraordinary twilight phenomena have been

well described by J. Kiessling.** One of these phenomena, the Bishop’s

ring, has been the subject of many investigations.t® The actinometric

°E. KwLuGe: “Ueber einige neue Forschungen auf dem Gebiete des Vulkanismus,”

Zeitsch. d. deut. geol. Ges., vol. 15, p. 377. 1863.

10 JOSEPH P. O’REILLY: “On the Dates of Volcanic Eruptions and Their Concordance

with the Sun-spot Period,’’ Proc. Roy Irish Acad., 3 ser., vol. 5, p. 392. 1899.

4 CH. V. ZENGER: “La théorie électrodynamique du monde et les éruptions volcaniques

et grands sismes,” Assoc. franc. p. l’avancement des sciences, Sess. 33, p. 572. 1904.

2H. I. JENSEN: “Possible Relation Between Sunspots and Volcanic and Seismic Phe-

nomena and Climate,” Jour. Roy. Soc. of New South Wales, vol. 38, p. 40. 1904.

13R. D. M. VERBEEK: Krakatau. Batavia, 1885.

“G. J.SyMons: The Eruption of Krakatoa and Subsequent Phenomena. London, 1888.

* J. KIESSLING: Untersuchungen tiber Diimmerungserscheinungen zur Erkliirung der

nach dem Krakatau-Ausbruch beobachteten atmosphirisch-optischen Stérung. Hamburg,

1888.

16 J. M. PERNTER: Meteorologische Optik, pp. 469, 769. Wien, 1902-10. See also —

SERENO BISHOP: ‘“‘The Origin of the Red Glows,’ Amer. Meteor. Jour., vol. 3, pp. 127,

193. 1886.

152 ANNALS NEW YORK ACADEMY OF SCIENCES

observations made by A. Crova,™? at Montpellier, have frequently been

cited as a proof of the decrease of solar radiation due to a purely ter-

restrial cause. Curiously enough, the effect of the Krakatoa dust veil

on atmospheric temperature seems to have attracted no special attention.

Besides the Krakatoa, other volcanoes were very active during the year

1883. Of these St. Augustin*® and Bogoslof*® of the Aleutian chain of

islands, as well as the Ometepe,*® may be mentioned.*t The violent

eruption of St. Augustin, which occurred on October 6, is of particular

interest. St. Augustin is an island south of the Alaskan peninsula

(position—153° 25’ W., 59° 18’ N.). The explosion split the island

in two, from peak to base, while the greater portion of the northern half

of the volcano was blown away.

The study of the temperature data for the year 1902 is also of special

interest, not only because during that year the world’s volcamic activity

was greatly intensified, but also because some of the explosive eruptions

which occurred undoubtedly produced a dust veil in the higher layers of

the atmosphere.

A decrease of solar radiation has been observed by Henri Dufour? in

Lausanne, by Harvey N. Davis”? in Providence, by H. H. Kimball?* in

North Carolina, by S. P. Langley?®? in Washington, by L. Gorczynski’*

in Warsaw, and this decrease has generally been ascribed to the presence

of. volcanic haze.

In 1901 the outbursts of Mt. Colima, Mexico, were more frequent and

more intense than during the preceding years. The same was true in

7A, Crova: “Sur les observations actinométriques faites 4 Montpellier,” Comptes

Rendus...vol. 106, p. 810. 1888.

18 GEORGE DAVIDSON: ‘“‘Notes on the Volcanic Eruption of Mount St. Augustin,’’ Science,

vol. 3, p.186. 1884.

19C, Hart MERRIAM: “Bogoslof, our Newest Volcano,’? Harriman Alaska Expedition.

Volo 2.1p21 291 LO0N

20G. MERCALLI: Vulcani attivi della terra, p. 356. Milano, 1907.

2C,. W. C. Fucus: ‘Die vulkanischen Ereignisse des Jahres 1883,’ Min. u. petrogr.

Mitth., vol. 6n. F., p. 185. 1885. ;

~ HENRI Durour: ‘Sur la diminution du rayonnement solaire,”’ Comptes rendus, vol.

SGnps clas) 19033

°3 HARVEY N. Davis: ‘Observations of Solar Radiation with the Angstrém Pyrheli-

ometer,’’ Monthly Weather Rev., vol. 31, p. 275. 1903.

2H. H. KIMBALL: ‘Observations of Solar Radiation with the Angstr6ém Pyrheliometer

at Asheville and Black Mountain, N. C.,’’ Monthly Weather Rev., vol. 31, p. 320. 1903.

See also same author: “Solar Radiation, Atmospheric Absorption and Sky Polariza-

tion, at Washington, D. C.,’”’ Bull. Mt. Weather Observatory, vol. 3, p. 110. 1910.

228. P. LANGLEY: “On a Possible Variation of the Solar Radiation and its Probable

Effect on Terrestrial Temperatures,’ Astroph. Jour., vol. 19, p. 305. 1904.

76 LADISLAS GORCZYNSKI: “Sur la diminution de l’intensité du rayonnement solaire en

1902 et 1903,” Compt. Rend. ... vol. 138, p. 255. 1904.

Same author: ‘“Quelques renseignements sur la dépression du rayonnement solaire i

Varsovie en 1903,” Bull. Météor. du départ. de l’Hérault. Montpellier, 1906.

ARCTOWSAKI, VOLCANIC DUST VEILS 153

1902 and even more so in 1903. The continuous observations of Mt.

Colima made at the Zapatlan Observatory** from 1893 to 1905 give, for

the years 1900 to 1904, the following frequencies of days during which

cruptions designated “erupcion grande” have been recorded: 15, 41, 37,

101, 2. According to C. G. Abbot,?* a photograph taken on March 7,

1903, shows a column of ashes reaching an altitude of about 17 miles.

‘his figure is a simple estimate and may be exaggerated.

On May 7, 1902, La Soufriére, St. Vincent, was in violent eruption.

“The particular feature of this eruption was the enormous amount of

dust which was thrown into the air and distributed over a vast, somewhat

elliptical area. . . . The British steamship ‘Coya’ had an eighth of

an inch of voleanic dust from this volcano fall on her deck when she was

two hundred seventy-five miles east-southeast of St. Vincent.” *° On

May 8, 1902, a sea of fire destroyed St. Pierre, Martinique. The follow-

ing violent eruptions of Mt. Pelé occurred on May 20 and 26, June 6,

July 9 and August 30.

The influence these eruptions may have had on the thermal trans-

parency of the higher atmospheric layers is questionable. The excellent

photographs taken by A. Lacroix*® show indeed that the occasional blasts

of incandescent gases and ashes did not exceed an altitude of 4,000 m.

Only an extremely small portion of the projected pulverized ashes must

have reached the average altitude of the cirrus clouds or even the strato-

sphere. This may not have been the case in the violent eruptions of the

Santa Maria volcano, in Guatemala. The eruption began on October 24,

1902. An eye-witness writes: “During the first four days of the eruption

no view could be had of the rising crater-cloud from the immediate

vicinity of the volcano. Only at a distance of forty miles to the north

and east could the erupted sand and smoke be seen against the sky.

‘wo days later I had another opportunity to view the eruption’ from a

distant hill under a clear sky, and in the day. The appearance then was

as follows: The peak of Santa Maria was sharply delineated against the

sky. To the westward or oceanward of this pyramid rose every few

minutes immense masses of globular clouds, like steam and smoke thrown

out of a locomotive when it first starts. The clouds rose to a height of

77S. Diaz: Efemerides dél volcan Colima. Mexico, 1906.

°C. G. ABBoT: ‘Do Volcanic Explosions Affect our Climate?’ Nat. Geog. Mag., vol. 24,

mists 1913:

JOSE MARIA ARREOLA: “Brief Notice of the Observations of Colima,” Jour. Geol., vol.

op ato. 1903:

* EDMUND OTIS Hovey: ‘Martinique and St. Vincent; a Preliminary Report upon the

Eruptions of 1902,” Bull. Am. Mus. Nat. Hist., vol. 26, p. 333. 1902.

50 A. LACROIX: La montagne Pelée et ses éruptions. Paris, 1904.

154. ANNALS NEW YORK ACADEMY OF SCIENCES

20,000 feet above the crater in three or four seconds.” . . .*1. The fall

of ashes was observed principally in a northwestern and northern direc-

tion. In Chicharras, on the Mexican border, the thickness of ashes that

fell on the ground was 420 mm., whereas in Oaxaca and San Juan

Bautista only 5 mm. were observed.*?

The eruption of the Mua, on Sawaii of the Samoa Islands, which oc-

curred October 30, 1902, was not violent enough to be taken into con-

sideration.** ‘The same may be said about the Isalco eruption, in Sal-

vador.** On the contrary, the Tori-shima eruption of August 7 and 9,

1902, seems to have been very violent. Reports of this eruption have

been published by F. Omori and others of the Imperial Earthquake In-

vestigation Committee, but these publications were not accessible to the

writer.

There can be no doubt that during the year 1902 a considerable quan-

tity of pulverized lava must have been projected into the higher layers

of the atmosphere, above the clouds. Bishop’s ring was observed anew,”°

as well as extraordinary twilight phenomena ;** but a comparison is hardly

possible with those which were due to the Krakatoa eruption. One single

volcanic explosion, if sufficiently violent, may therefore obscure the

stratosphere very much more than a score of violent eruptions of a less

explosive character.

' This seems to have been the case of the Katmai eruption. “Katmai

volcano is in the Aleutian Range, Alaska, latitude 58° N., longitude 155°

W. approximately. On the afternoon of June 6, 1912, it suddenly be-

came explosively eruptive, continued in a state of great activity for about

three days, and was reported to be still somewhat active at the end of

October, 1912.” ** For particulars I will refer to an account published

31 GUSTAV BISEN: ‘The Earthquake and Volcanic Eruption in Guatemala in 1902,”

Bull. Am. Geog. Soc., vol. 35, p. 391. 1903.

22 KarL SAPPER: “‘Die vulcanischen Ereignisse in Mittelamerika im Jahre 1902,” N.

Jahrb. f. Miner., Jahrg., 1904, vol. 1, p. 39. See also ALFRED BERGRAT: ‘‘Die Produkte

der letzten Eruption am Vulkan §S. Maria in Guatemala,” Centr. f. Mineral., Jahrg.

1903, p. 112.

33 GEORG WEGENER: ‘“‘Die yvulkanischen Ausbriiche auf Sawaii,” Zeit. Gesell. f. Erdkunde

zu Berlin, p. 208. 1908.

3K aRL SAPPER: “Die jiingsten Ereignisse am Vulkan Izalco,” Centr. f. Mineral..

Jahrg. 1903, p. 103. 1908.

35. A. ForEL: ‘“‘Le cercle de Bishop de la Montagne Pelée 1902-1904,” Arch. Se. Phys.

et Nat., Ser. 4, vol. 19, p. 229. 1905.

36 P. GRUNER: ‘‘Ueber die neuen Dimmerungserscheinungen,” Mitt. naturf. Gesell. Bern.

a. d. J. 1903, p.1. 1904.

E. MARCHAND: “Les lueurs crépusculaires et phénoménes connexes,” Annuaire Soc.

Météor. France, vol. 53, p. 40. 1905.

87 HERBERT H. KIMBALL: “The Effect upon Atmospheric Transparency of the Eruption

of Katmai Volcano,’ Month. Weather Rev., vol. 41, p. 153. 1913.

ARCTOWSKI, VOLCANIC DUST VEILS 155

in the National Geographic Magazine.*® A summary of the effects of

the Katmai eruption on atmospheric optical phenomena has been given

by J. Maurer and C. Dorno.*®

The fact that the Katmai eruption occurred in a far northern latitude,

and has not been followed by similar volcanic outbreaks in other parts

of the world, is most valuable. Since the general atmospheric circulation

of the southern hemisphere is independent of that of the northern hemi-

sphere, it is difficult to imagine how the haze produced by the Katmai

eruption could have been carried south of the equator. If therefore we

observe, on the temperature curves of stations belonging to the southern

hemisphere, great similarities in the details of the curves of stations

belonging to the northern hemisphere, attributed to the presence of vol-

canic haze, it is evident that, however incomplete our knowledge of the

general atmospheric circulation may be considered, the supposed volcanic

dust influence must be discarded.

Some argumentation, however, leaves the question open for discussion.

The correlations between the seasonal variations of far distant stations,

first noticed by H. W. Dove*®, then later more extensively studied by

H. F. Blanford,** H. H. Hildebrandsson,*? J. Hann,** H. G. Lyons,**

Felix M. Exner,*? R. C. Mossman*® and others, show that one may pre-

_ sume that an anomaly at one “center of action” of atmospheric circula-

tion of one hemisphere will produce a similar anomaly at a corresponding

station belonging to a correlated “center of action” of the other hemi-

sphere. I suppose that the number of stations for which temperature

records have been compared is sufficient to eliminate the possibility of

such an argument.

33 GEORGE C. MaArTIN: ‘The Recent Eruption of Katmai Volcano in Alaska,” Nat,

Geog. Mag., vol. 24, p. 131. 19153.

% J. MAURER and C. Dorno: ‘Ueber den Verlauf und die geographische Verbreitung

der atmospherisch-optischen Stérung 1912-1913,” Meteor. Zeit., vol. 31, p. 49. 1914.

40H. W. Dove: Nicht periodische Verinderungen der Verbreitung der Wiirme auf der

Erdoberfliche. Berlin, 1869.

41H. F. BLANForD: “On the Barometric See-saw between Russia and India in the

Sun-spot Cycle,’’ Nature, vol. 21, p. 477. 1880.

#H. H. HILDEBRANDSSON: “Quelques recherches sur les centres d’action de l’atmos-

phere,” K. Svenska Vetensk. Akad. Handlingar, vol. 29, no. 3, vol. 32, no. 4, vol. 45,

no. 2, vol. 45, no. 11. 1897, 1899, 1909, 1910.

4 J. HANN: “Die Anomalien der Witterung auf Island in dem Zeitraume 1851-1900

und deren Beziehungen zu den gleichzeitigen Witterungsanomalien in Nordwesteuropa,”’

Sitz. Math. nat. Kl. K. Akad. d. Wiss., vol. 113, II a, p. 183. Wien, 1904.

4*H. G. LYONS: The Physiography of the River Nile and its Basin. Cairo, 1906.

4 FELIX M. EXNER: ‘‘Ueber monatliche Witterungsanomalien auf der nérdlichen Ird-

halfte im Winter,” Sitz. K. Akad. Wiss. Math.-Naturw. Klasse., vol. 122, II a, p. 1165.

Wien, 1913.

_ 4#R. C. Mossman: “Southern Hemisphere Seasonal Correlations,’’ Symons’s Met. Mag.,

vol. 48. London, 1913.

156 ANNALS NEW YORK ACADEMY OF SCIENCES

PIKE’S PEAK TEMPERATURE RECORDS

The meteorological observations made on the summit of Pike’s Peak

extend through the years 1874-1887. It seems to me that the records

of this station—situated near the center of the North American conti-

nent, at an altitude of 14,111 feet—may be considered as being most

reliable material for the study of the influence of the dust veil of the

years 1883 and 1884, upon temperature conditions in the United States.

Table I gives the recorded mean monthly temperatures (5 A. M., 1, 9

P. M.) expressed in departures from the corresponding monthly means

of the entire series of observations.** In Figure 1 the consecutive twelve

monthly means are represented graphically. On this diagram, one notices

Fic. 1.—Curve of the consecutive means of temperature observed on the summit of

Pike’s Peak

immediately that the curve is abnormal between the crests C and F, that

in order to have a more regular variation, we ought to have a crest (D)

in place of the depression K. Admitting the existence of such a crest,

we have intervals of 33, 27, about 31 and about 35 months between the

crests and 30, 33, 25 and 31 months between the depressions. These

tigures are similar to those obtained from the records of many other

stations. But, as on the curves expressing the succession of consecutive

means for longer series of observations—for example those of Batavia,

New York, Rome, Warsaw and Stockholm, which are at present at my

disposal—the pleionian crests do not always reoccur at approximately

regular intervals and seem to be missing sometimes, just like the pre-

sumed crest (D), the fact that the Pike’s Peak curve shows a long in-

47 Annals Astron. Obs. Harvard College, vol. 22, p. 457. 1889.

157

ARCTOWSKI, VOLCANIC DUST VEILS

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158 ANNALS NEW YORK ACADEMY OF SCIENCES

terval between the crests C and £, of nearly six years or the double of

the interval generally observed, cannot be considered as an argument in

favor of a correlation between the temperature depression K and the

eruptions of Krakatoa and St. Augustin. But the drop of temperature

for the mean of November, 1882,-October, 1883, comcides with the ap-

pearance of the Krakatoa veil in the States and also with a possible

influence of the Si. Augustin eruption, so that a simple chance circum-

stance can hardly be admitted. Then the curve starts up, just as it went

up before the consecutive mean ending with October, 1883, and this

tendency is maintained for a few months. The small crest d in the

depression K seems to be another detail showing clearly that this de-

nression is abnormal, that it is not a true antipleionian depression.

Besides, on the table of monthly departures, we notice the exceptional

figure—5.1 °F. for October, 1883, and the succession of negative de-

partures for the year 1884.

In order to show that the depression K was really abnormal and that

under normal conditions we ought to have had a pleionian crest (D) in

place of this depression, I will consider now the consecutive temperature

curve of Port Darwin, a station of North Australia, 12° 28’ lat. S.

TEMPERATURES AT PorRT DARWIN AND SOME OTHER STATIONS

In Table II the monthly temperatures for the years 1880-89 are given

in departures from the means figuring on top of the table.*® The dia-

gram (Fig. 2) represents the succession of consecutive annual means.

Fie. 2.—Thermopleionian variation of Port Darwin

A comparison of Figures 1 and 2 shows striking similarities between

the two curves. IJ marked, therefore, the crests of the Port Darwin curve

#8 Meteor. Observat. made at the Adelaide Observatory...during the year 1880...1889.

159

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tite eerezg

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160 ANNALS NEW YORK ACADEMY OF SCIENCES

with the letters C', (D), & and F’, expressing that way my view that the

variations are identical.

However, a very important difference must be noted at once; it is the

difference in time of the occurrence of the same pleionian crests and

antipleionian depressions. The Pike’s Peak maxima occur ten or eleven

months later than those of the Port Darwin curve. Now, the distance

between the crests C and # is in both cases 67 months. Some of the

details of the crest C of Pike’s Peak, as well as of the depression that

followed, may be easily observed on the corresponding crest and depres-

sion of the Port Darwin curve. If, therefore, the missing pleion of the

Pike’s Peak curve appears very plainly on the Port Darwin curve, as it

does, we are justified in presuming that the interval of more than 5 years

separating the crests C and # on the Pike’s Peak was indeed abnormal,

as well as the depression K, and that this anomaly must be ascribed to

the Krakatoa dust veil.

The crest (D) of Port Darwin is not developed to its normal value.

On the diagram the dotted line indicates the portion which must be con-

sidered as having been cut away. The anomaly begins with the consecu-

tive mean of October, 1882, to September, 1883, that is to say, just one

month sooner than on Pike’s Peak. This anomaly is not ten or eleven

months in advance on the corresponding detail of the Pike’s Peak curve,

but just one month and occurs one month later than the Krakatoa erup-

tion—when the dust veil reached Port Darwin, after having traveled

twice around the world along the equator. The duration of the anomaly

extends from the mean of October, 1882,-September, 1883, till the mean

of February, 1884,-January, 1885, or 17 months. The same figure may

be adopted in the case of the Pike’s Peak curve.

An important question arises now. Is it possible to estimate the lower-

ing of temperature due to the presence of the Krakatoa dust veil? Ac-

cording to the dotted lines of the diagrams the lowering of the tempera-

ture for the consecutive mean of September, 1883,-August, 1884, may

have been 1.9 °F. in the case of Port Darwin and 3.4 °F. in the case of

the Pike’s Peak observations. But this is an estimate of no scientific

value. ‘The departures of Tables I and II are of no help. It would be

necessary to know what these departures ought to have been. I imagine

it would be possible to attempt the calculations by tracing maps and by

comparison of the temperature conditions of the dust-affected regions

with those over which the dust veil was not spread out. But even in that

case comparisons would be most difficult, because we do not know how

the dust veil affected the general atmospheric circulation or how the

abnormal conditions of one region affected the temperatures of other

regions mechanically.

ARCTOWSKI, VOLCANIC DUST VEILS 161

The curve of the consecutive means of the temperatures observed at

the Batavia observatory confirms the results obtained so far. So do the

curves of Singapore, Port Blair, Colombo, Bombay and Aden,*® which

are reproduced in Figure 3.

The curves of Bombay and Port Blair show distinctly the antipleionian

depressions preceding and following the abraded pleionian crest. At

Singapore

Pek Blair.

Colombo.

Romboy :

Fic. 3.—Temperatures recorded at some Asiatic stations

Port Blair the pleion is less depressed; at Aden the most. If the lowest

mean of the two years following the Krakatoa eruption expresses the

maximum effect of the dust veil, we must say that this maximum effect

was not simultaneous at the different stations taken into consideration.

The following tabulation gives the periods, as well as the lowering of the

mean below the value of the last unaffected consecutive mean.

42 Singapore from: ‘Met. obs. at the foreign and colonial stations of the Royal Engi-

neers. ..1852-1886.’’ London, 1890.

The other stations from: ‘‘Report on the meteorology of India in 1880. ..1890.”

162 ANNALS NEW YORK ACADEMY OF SCIENCES

Pike’s Peak. Sept. 1883-Aug 1884:10°F below the mean of Sept. !882-Aug. 1883

Port Blair... Dec. 1883-Nov. 1884: 1.4 a ss a a a ** =

Bombay .... June 1883—May 1854 :0 6 ce os y e e i

Singapore... Sept. 1883-Aug 1884:0 6 im s ie °F = ce

Agden 2 ae... July 1883-June 1884 : 0.7 de “ a af a “e ee

Colombo.... Noy. 1883-Oct. 18584: 0.7 Ss o oe is om . eS

Port Blair... Nov. 1883-Oct. 1884:0 2 a “ = ee ‘iy

TEMPERATURES AT FORT-DE-FRANCE, MARTINIQUE

During the terrific eruptions of Mt. Pelé, on May 8 and 20, 1902, the

usual meteorological observations were made at Fort-de-France. In

Table III the monthly mean temperatures are given in form of departures -

from the corresponding means of the years 1900-1909 and in Figure 4

ihe curve A represents these departures graphically. Curve B shows the

Mires. ]

eee |

| pee I

i 1 | X

| | i

Fic. 4.—Temperature records at Fort-de-France

oie ee

succession of consecutive yearly means and C gives the values of the

aifferences: between the monthly mean maxima and minima. This last

curve indicates the changes of the range of the diurnal variation of

temperature. On the average, the lowest value is observed in July and

the highest in March. The respective mean values for the different

months are: 8°.32, 8°.72, 8°.95,-8°.72; 7°.98, 7°.18)6°.99, 7° 5am ee

163

ARCTOWSKI, VOLCANIC DUST VEILS

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164 ANNALS: NEW YORK ACADEMY OF SCIENCES

8°.14, 8°.17 and 8°.02 C. Curve C indicates a long range variation

completely independent of the temperature variation, which has a maxi-

mum in 1902 and a minimum in 1910, as can be seen on curves A and

B. The data which I collected some years ago from the observations of

Russian stations®® give the result that these long-range variations of the

values of the mean diurnal oscillations of temperature differ from one

region to another and that these variations correspond to changes of

cloudiness. An increase of cloudiness diminishes the average daily range

of temperature. Considering now the trend of the curve we notice a

more or less progressive increase of the values from 1901 to 1904. This

is just the contrary of what would be expected. At Fort-de-France the

daily range of temperature has therefore not been greatly affected by the

dust veils produced by the eruptions of Mt. Pelé, Sta. Maria and Mt.

Colima.

The mean temperatures have also been affected but very slightly. The

pleionian crest of 1902-1903, as indicated on the curve of consecutive

means, has been depressed a little, but certainly not more than 0.15 °C.

or 0.2 °F. It is difficult to judge how much the mean temperatures of

the individual months have been affected. Curve A shows a decrease of

the departures for the months of June, July and August, 1902, following

the great Mt. Pelé eruption: also a more pronounced decrease for Novem-

ber and December, possibly due to the Sta. Maria eruption, and finally

the low April departure may have been caused by the Colima eruptions.

However the departures of the months of May, 1902, to the end of 1903

are all above the average and if the slight deflections observed during the

period of great volcanic eruptions must really be attributed to dust veils,

it may be presumed that the means of some months have been affected

more than those of other months but none sufficiently to mask the plei-

onian character of the departures. Moreover, the effect of the dust veil

ceased long before the complete development of the antipleionian de-

pression of 1904-1905. This antipleion cannot, therefore, be considered

as being a consequence of the formation of the volcanic dust veil.

TEMPERATURES AT PARA, CAYENNE AND THE WEST INDIES

Although the different examples I have given may be considered as a

sufficient proof of the fact that the pleionian variations are absolutely

distinct in their origin from the possible temperature variations due to

the presence of volcanic dust, I will examine a few more consecutive tem-

50 HENRYK ARCTOWSKI: “Notice sur les yariations de longue durée des amplitudes

moyennes de la marche diurne de la température en Russie,’ Bull. Soc. belge d’Astrono-

mie. Bruxelles, 1908.

ARCTOWSKI, VOLCANIC DUST VEILS ta5

perature curves simply to show how cautious one has to be in the study

of the cause of local climatic variations.

In the following diagram (Fig. 5), I reproduce the curves of the con-

secutive means of temperature for Para,°* Cayenne** and the West Indian

a p ob i) *]

ans

Fic. 5.—Comparison of the consecutive temperature curves of Cayenne and Paré with

those of the West Indies

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stations: Port-au-Prince,** St. Croix Christianssted,®+ St. Lucia and

Barbados,®* together with the curve of Fort-de-France.

The curves of St. Lucia and Barbados are so very different from the

curve of Fort-de-France that one would be inclined to admit some errors

1 Meteorologische Zeitschrift, vols. 23, 24, 28, 31, pp. 517, 431, 215, 139.

52 Annales Bur. Centr. Météorol. de France.

*S Annales Bur. Centr. Météorol. de France.

** Meteorologisk Aarbog for 1900...1909. Udgivet af det danske meteor. Institut.

* Received in manuscript from ihe Meteorological Office in London.

166 ANNALS NEW YORK ACADEMY OF SCIENCES

of observations. But the curve of St. Lucia is a transitional form be-

tween those of Fort-de-France and Barbados, and Para, in Brazil, 1°

27’ S. lat., shows, between 1902 and 1905, exactly the same abnormal

depression as Barbados. A very accentuated depression between 1903 and

1904 is also ‘characteristic for Arequipa and Maguritius®® as well as St.

Helena.*’ The temperature curve of Apia, Samoa Island, displays the

same very pronounced antipleionian depression, corresponding perfectly

with that of Barbados. It is difficult, therefore, to avoid the conclusion

that the curve of Barbados must be correct and that if it differs very

greatly from the curves of the other West Indian stations this fact may

simply be considered as an interesting subject for special investigation.

For my present purpose the Barbados curve is a very valuable argu-

ment against a possible supposition in favor of the voleanic dust hypoth-

esis. One could imagine that if the Fort-de-France temperature records

as well as those of some other stations of the West Indies have not been

very much affécted by the eruption of Mt. Pelé and the other volcanoes,

it may be because the dust veils have been carried into the southern

hemisphere, and that there the curves of the stations belonging to the

belt of the southern trade-winds show the depression just as much, if not

more so, than in the case of the Krakatoa eruption.

The fact that Cayenne, which lies between Para and Barbados, has a

curve resembling those of St. Croix and Port-au-Prince and that these

stations belong to the northern trade winds, and the fact that the curve

of Barbados is similar to: those of the southern stations, show that this

explanation of the observed anomaly is inadequate.

TEMPERATURES OBSERVED IN ALASKA

Assuming that the voleanic haze produced by the Katmai eruption of

June 6, 1912, must have had the greatest effect on the temperatures re-

corded in Alaska and in Canada, I will now compare the curves of seven

stations in Alaska with the curves of Victoria and Edmonton, Mauritius

and Arequipa.

_The temperature data for the Alaskan stations were obtained from the

United States Weather Bureau, the Arequipa daily observations were

sent to me by Professor Edw. C. Pickering and the Mauritius data by

Dr. H. R. Mill and by the director of the Mauritius observatory. The

observations considered extend over the five years 1909-1913.

Since Mt. Katmai could not have affected the temperature conditions

of Arequipa and Mauritius, it is safe to take the curves of these stations

56 ARCTOWSKI: Op. cit.. Bull. .Am..Geog. Soc., vol. 44, p. 598.

57 ARCTOWSKI: Op. cit. Annals N. Y. Acad. Sci., vol. 24, p. 109. 1914.

ARCTOWSKI, VOLCANIC DUST: VEILS 167

as a standard. Moreover, in my previous publications I have shown that

the consecutive means observed at Arequipa express very well the normal

pleionian variation and may serve as a standard in all cases of comparison.

In the accompanying diagram (Fig. 6) the consecutive temperature

curves along the Pacific coast are represented, together with the Mauritius

and Arequipa curves. The latitudes

of Fort Liscum, Sitka, Loring and

Maetorta’ are: 61° 2’, 56°50", 55°

32’ and 48° 26’ N.

The occurrence of the eruption

coincided with the pleionian crest of

Arequipa. For Arequipa the con-

secutive mean of July, 1911, to

June, 1912, is the highest. From

then on the temperature decreases

until the consecutive mean of Octo-

ber, 1912,-September, 1913. The

same is true at Mauritius; but there

the pleionian crest is very different

from what it is at Arequipa—it is

flat. Since the same may be ob-

served on the curve of Victoria, the

apparent depression of the crest of

this station cannot be ascribed with-

out hesitation to the effect of the

Katmai haze. ;

The other curves—the Fort Lis-

cum curve in particular—resemble

the Arequipa curve so very much

that we may admit that at least the

depression of the values from the

mean of August, 1911,-July, 1912,

to the mean of December, 1911,-No-

vember, 1912, is due to the effect of voleanic dust. The dotted lines

indicate the portions of the pleionian crests which may have been de-

pressed. : vas |

_ The curves of the stations at Nome, Tanana and Fairbanks, situated

north of the Alaskan range, do not seem to have been affected, and that

of Eagle but slightly. In the case of Edmonton, it is difficult to decide.

i): 1910 1g gy 1g'3.

Avequina.

Fort Lligcum .

Sitka -

Loring.

Nedestée ’

he Mauriti ws.

Rx equs pa: |

Fic. 6.—Arequipa pleion of 1911-1912

observed in Alaska

168 ANNALS NEW YORK ACADEMY OF SCIENCES

TEMPERATURE VARIATIONS AT STOCKHOLM AND BATAVIA

In a recent publication,** Axel Wallen gives the curve of the consecu-

tive annual means of temperature observed at Stockholm, for the entire

series of observations extending from 1756. A comparison of his curve

with the Batavia curve which I published some years ago shows that they

are very much alike. The monthly departures of Stockholm*® present

also many striking analogies with those of Batavia.°° It would be worth

while to make a more detailed examination.

The variations at Stockholm and Batavia seem to be in perfect har-

mony: a crest of the Stockholm curve corresponds nearly to each crest of

the Batavia curve, and the intervals between corresponding crests are

approximately the same. It will be necessary to find many more such

similar curves, belonging to far distant stations, before the main ccn-

clusion which might be inferred from the comparison of the Batavia and

Stockholm curves will have the practical value one might expect.

Batavia being in advance of Stockholm, it is easy to understand that,

taking seasonal departures—as Hildebrandsson, Mossman and others have

done—correlations between these two stations have to be found. But

these correlations will evidently be simply apparent correlations. It is

not because a given season of the year is abnormally warm in Batavia,

that the following season has to be exceptionally warm in Stockholm and

so on; but it is because the same pleionian variation appears with the

same regularity in one place as in the other, but slightly retarded, that a

seasonal correlation between the two places must exist. The Batavia and

Stockholm curves give, therefore, a plausible explanation to the so far

unexplained correlations between seasonal departures of distant stations.

Coming back to the volcanic dust problem studied in this paper, it is

important to notice that the Stockholm curve seems to be more complete

than the Batavia curve. The pleion of 1883, which on the Batavia curve

has been greatly reduced by the dust veil effect, is very much better

developed on the Stockholm curve. In a similar way some apparent

anomalies on the Batavia curve may be considered as being really anom-

alies which would not exist if the curve were perfectly developed as it

ought to be. For example, the portion of the consecutive temperature

curve of Batavia from 1871 to 1873 is abnormal. Likewise, the long

°S AXEL WALLEN: “Flerariga variationer hos vattenstandet i Milaren...,’ Meddelan-

den fran Hydrografiska Byran: 4. Stockholm, 1913.

5° H. E. HAMBERG: ‘‘Moyennes mensuelles et annuelles de la temperature...a l’obser-

vatoire de Stockholm,’ Konigl. Svenska Vetenskaps-acad. Handlingar, vol. 40, no. 1.

Stockholm, 1906.

8° Observations made at the Royal Magnetical and Meteorological Observatory at Bata-

via. Vol. 28, p.108. 1907.

ARCTOWSKI, VOLCANIC DUST VEILS 169

interval between the pleionian crests of 1891 and 1897 of the Batavia

curve is due to a missing crest which is well marked on the Stockholm

curve.

SUN-SPOTS AND ATMOSPHERIC T'EMPERATURE

The conclusion so far is that the thermopleionian variations cannot be

ascribed to more or less frequent reoccurrences of volcanic dust veils. In

exceptional cases, such as the Krakatoa eruption, atmospheric tempera-

ture has been affected practically all over the world; but even then the

pleionian crests of the consecutive yearly temperature curves have simply

been partially erased, or modified in such a way that there can be no

doubt that the antipleionian depressions which followed were not due to

the presence of volcanic haze.

The hypothesis that the pleionian variations of temperature have an

extra-terrestrial cause is therefore a perfectly logical conclusion.

In order to prove that this hypothesis is well founded, it will be neces-

sary to demonstrate the existence of a correlation between atmospheric

temperature and solar radiation, or perhaps some other phenomenon of

which the “solar constant” is a function.

It has been presumed that differences in the extent or relative position

of the Zodiacal light might be such a phenomenon.®* Variations of the

solar corona could possibly also have some effect on the radiant energy

transmitted towards the earth ;°* but it seems more probable that atmos-

pheric temperature is directly affected by changes occurring on the sun’s

surface or solar atmosphere.** The well known sun-spot cycle of about

11 years’ mean duration proves it very well.

It was Alfred Gautier®* who, as far back as 1844, discovered the rela-

tion that exists between atmospheric temperature and the solar cycle.

6. BELOT: “‘La matiére zodiacale et la constante solaire,”’ C. R. Ac. Sci. Paris, vol.

£59, p. 157. 1913.

8 A. Crova: “Etude de l’intensité calorifique de la radiation solaire au moyen de

lactinométre enregistreur,’’ Annales de chimie...ser. 6, vol. 14, p. 543. Paris, 1888.

Some arguments in favor of this hypothesis may also be found in the following

papers:

S. P. Lanciey: ‘‘The Heat Radiation of the Corona,’ Astroph. Journal, vol. 12, p.

aor 1900:

C. G. Apsot: “‘A Bolometric Study of the Solar Corona,’ Publ. Astron. Soc. Pacific,

vol. 20, p. 86. 1908.

JOHN A. MILLER: “The Position of Certain Coronal Streams on the Assumption

that the Corona is a Mechanical Product,” Astroph. Jour., vol. 33, p. 303. 1911.

8S. P. LANGLEY: “The Solar Atmosphere, an Introduction to an Account of Researches

made at the Allegheny Observatory,’’ Amer. Jour. Sci., ser. 3, vel. 10, p. 489. 1875.

J. HAHN: “Contributions to the Theory of the Sun,’ Annals Roy. Obs. Edinburgh,

vo. 1, p. 74. 1902.

6 ALFRED GAUTIER: “‘Recherches relatives 4 l’influence que le nombre et la permanence

des taches observées sur le disque du soleil peuvent exercer sur les temperatures terres-

tres,’ Annales de chim. et de phys., ser. 3, vol. 12, p. 57. 1844.

170 ANNALS NEW YORK ACADEMY OF SCIENCES

Since then many researches have been made. A recent publication by

W. Koeppen® gives the most convincing data. Koeppen utilized the

annual temperature means collected by J. Mielke,®® as well as the figures

which served him in previous publications on this subject, and completed

the tables in order to have the results of practically all available obser-

vations made, all over the world, from 1811 to 1910. The final result

may be advantageously reproduced in a form shghtly different from that

given by Koeppen.’

The following figures represent the mean departures from the general

means for the years of sun-spot maxima and of those before and after

these years:

ED ONG ae

0.45 | -0.15 | -Yovonee!

+0.09 | —0.03 _—0.15 | —0,28

The means of the years of sun-spot minima, as well as the means of

the years preceding and following these minima, are given below:

Tae Se —1

+-O td.

—0.11| +0.07 | +0.24

+2 +3 years

+0.04

+0.22 0.00 °C.

The conclusion is that the solar cycle of about 11 years’ mean duration

has a well pronounced influence upon atmospheric temperature, and that

the difference between maxima and minima equals on the average nearly

Qian!

But the diagram representing the succession of the preceding figures

graphically shows that the 11-year period 1s complicated by the existence

of a shorter variation of less importance. It seems that during the sun-

spot cycle, temperature undergoes a double oscillation; that between the

consecutive temperature minima, corresponding to sun-spot maxima,

there are two minima and three maxima, one of which corresponds to the

sun-spot minimum. The diagram given by Koeppen for the Russian —

temperature data is perhaps the best illustration of this fact.

8 W. KOoOEPPEN: “‘Lufttemperaturen, Sonnenflecke und Vulkanausbriiche,”’ Meteor.

Zeitsch., vol. 31, p. 305. 1914.

66 JOHANNES MIELKE: “Die Temperaturschwankungen 1870-1910 in ihrem Verhdltnis

zu der 11 jaihrigen Sonnenfleckenperiode,’’ Aus dem Archiv der deutschen Seewarte, vol.

36;-N0) Oo. LOS: :

87 Koeppen adds the annual temperature departures of the years. of sun-spot minimum,

one year after, two a. s. f. This gives him 11 columns of figures. The disadvantage of

this method is that the year of sun-spot maximum occurs once in column 4, 3 times in

column 5, 4 times in column 6 and once in column 9. His first column of figures com-

prises the departures for the years 1811, 1822, 1833, 1844, 1855, 1867, 1878, 1889 and

1900. Only four of these years are years of sun-spot minima; the others precede or

follow such a year. However it would have been difficult to make a better adjustment

than Koeppen did.

ARCTOWSKI, VOLCANIC DUST VEILS 41

In the case of rainfall, Charles Meldrum®* long ago noticed the double

oscillation. For the weather conditions in the United States, Frank H.

Bigelow® found that “the occurrence of four subordinate crests in the

11-year periods suggests strongly that a 234-year period is superposed

upon the long sweep of that periodic curve.” Later, Bigelow’ discovered

a correlation between this short period variation and the frequency

changes of solar prominences.

One may conclude that Koeppen’s calculations give only a first ap-

proximation, and that in reality the phenomenon of the 11 years’ varia-

tion is complicated by a variation of a shorter periodicity, perhaps also

in connection with the changes of sun-spot frequency.

A strong argument in favor of this supposition is the circumstance

that the commonly known 11 years’ variation is not the only sun-spot

eyele. The relative numbers of Wolf and Wolfer have been analyzed

by Arthur Schuster,"* who found the following periods: 13.5, 11.125,

8.36, 5.56, 4.79, 3.71. Hisashi Kimura,’? on the other hand, found

periods of 16.59, 13.53, 11.114, 9.99, 8.25, 5.95, 5.49, 4.85 and 3.63 years.

According to the most recent investigations** it may be, however, that,

“while these various periods are apparent, yet many are illusory.”

All these researches show very plainly that the frequency variation of

sun-spots is a most complicated phenomenon. So are also the tempera-

ture variations. It is certain that the pleionian variation of the Arequipa

or equatorial type has apparently nothing in common with the 11-year

cycle; but some correlations exist. In 1893 the antipleions were pre-

dominant. ‘The year 1900 was a pleionian year. One may presume,

therefore, that the solar cycle of 11 years has a direct influence on the

long range or macropleionian variations and that the shorter pleionian

and antipleionian oscillations of terrestrial temperature are the product

of some minor sun-spot cycle, or perhaps of some other solar cycle which

does manifest itself with the same evidence as the sun-spot frequency

variations.

As soon as the particulars of the pleionian variations are better known,

68 CHARLES MELDRUM: “Report on Sun-spots and Rainfall,’ Rep. 48th meeting Brit.

ASSOC, p. 257. 1879.

6 FRANK H. BiGELow: “Abstract of a Report on Solar and Terrestrial Magnetism in

their Relations to Meteorology,’ Bull. no. 21, U. S. Weather Bur., p. 125. 1898.

7 FRANK H. BicELow: “The Relations Between the Meteorological Elements of the

United States and the Solar Radiation,’’ Amer. Jour. Sci., ser. 4, vol. 25. p. 423. 1908.

71 ARTHUR ScHUSTER: “On the Periodicities of Sun-spots,’’ Phil. Trans. Roy. Soc.

London, vol. 206A., p. 69. 1906.

7 AHISASHI KriMuRA: “On the Harmonic Analysis of Sun-spot Relative Numbers,”

Monthly notices Roy. Astr. Soc., vol. 73, p. 545. 1913.

% A. E. DouGLass: “A Photographic Periodogram of the Sun-spot Numbers,’’ Astroph.

Jour., vol. 40, p.. 326. 1914.

172 ANNALS NEW YORK ACADEMY OF SCIENCES

it will be possible to reverse the problem and search in the variable solar

phenomena for the periodicities which might correspond to the changes

of atmospheric temperature. But the problem is extremely complicated.

All we know about climatic variations is derived from observations taken

at the earth’s surface. Now, Charles J. Kullmer** “has shown that in

the belt of the northern United States and southern Canada where storms

cn the average are most numerous, the number of storms varies almost

directly in harmony with the number of sun-spots, just as is the case

with tropical hurricanes. In other areas, however, the reverse appears

to be true, and there is a decrease in storminess. The general conclusion

seems to be that when sun-spots are few in number cyclonic storms move

in a great variety of tracks, but when spots are numerous the storms tend

to confine themselves to a few well-defined tracks, so that stormimess is

more or less restricted to certain areas within which it is highly concen-

trated.” ** If we bear in mind the fact that storms are centered by

columns of ascending air, we must presume that on the average differ-

ences in storm frequency (and intensity) must produce corresponding

differences in the vertical distribution of temperature. Ellsworth Hunt-

ington supposes that “in this way increased activity of the sun might

cause increased activity upon the earth, and the earth’s surface might

become cooler while the upper air above the reach of convection might

become warmer.” *® ‘This most interesting suggestion could be verified,

T suppose.

Utilizing the table of Leo Kelley, giving the annual frequencies of

eruptions,‘* as well as Koeppen’s worlds mean temperature table, I made

for both overlapping ten-yearly totals. The following diagram (Fig. 7)

represents the figures graphically and shows plainly that some sort of a

relation between the numbers of volcanic eruptions and atmospheric tem-

perature is undeniable. |

Temperature seems to be higher when the eruptions are more frequent

and when volcanic activity is attenuated, atmospheric temperature seems

also to be lower than the average. This result is not in favor of the

volcanic dust explanation of climatic variations and if taken in con-

nection with the remarkable coincidences between the well pronounced

** CHARLES J. KULLMER: “The Shift of the Storm Track,” In: E. Huntington: “The

Climatic Factor...*° p. 193. Carnegie Inst. Pub. No. 192. 1904.

® ELLSWORTH HUNTINGTON: Op. cit., p. 253.

*6 Manuscript communication. Huntington discusses the question at length in his

recent paper, “The Solar Hypothesis of Climatic Changes,’ Bull. Geol. Soc. Amer., vol.

2: p41 6.) FAI:

™ JOHN MILNE: “Eighteenth Report of the Committee on Seismological Investiga-

tions,” p. 65. Report of the S3rd meeting oi the Brit. Assoc. Birmingham. 1913.

ARCTOWSKI, VOLCANIC DUST VEILS ie)

pleions of the years 1883, 1902 and 1912 and the exceptionally violent

volcanic eruptions which occurred simultaneously, shows that it may be

that the volcanic agencies are affected by the same extra-terrestrial factors

which influence climatic conditions.

How this could be, is a most interesting question leading to many

investigations. One may presume, for example, some correlations be-

tween the pleionian variation and storm frequency and storm intensity

= | ail i al 60 | alt bo | ‘SAi+ Igec.

VE.

\BhI-to (B44. fo ¥Li-J0 I8B\- Go

Fig. 7.—World’s temperature and the variation of the frequency of volcanic eruptions

and then a correlation between the more or less accentuated rapid oscilla-

tions of atmospheric pressure and volcanic eruptions.

A relationship between the mean values of interdiurnal differences of

atmospheric pressure and the solar cycle has been discovered by R.

Merecki,’* and, on the other hand, the much discussed relation between

barometric changes from day to day, and the frequency of seismic and

volcanic phenomena may be accepted as an established fact.‘* One may

73R. MBPRECKI: “Wplyw zmiennej dzialalnosci slonca na ceczynniki meteorologiczne

ziemske,’”’ Prace mat.-fizyczne, vol. 19, p. 181. Warszawa, 1908.

7 F. OMorRI: ‘‘Note on the Long-period Variations of the Atmospheric Pressure,” Bull.

Imp. Earthg. Invest. Comm., vol. 2, p. 215. Tokyo, 1908.

H. J. JOHNSTON-LAvis: ‘‘The Relationship of the Activity of Vesuvius to Certain

Meteorological and Astronomical Phenomena,’ Proc. Roy. Soc. London, vol. 40, p. 248.

1886,

174 ANNALS NEW YORK ACADEMY OF SCIENCES

ask, therefore, if the amplitudes of the barometric oscillations vary in

harmony with the pleionian cycles.

CONCLUSIONS

). The hypothesis attributing most climatic variations to volcanic

st veils, although verified in some cases, presents more a purely theoret-

ical than practical interest.

2). The dust veil produced by the Krakatoa eruption affected atmos-

pheric temperature very greatly. ‘The violent volcanic eruptions of 1902,

as well as the Katmai eruption of 1912, influenced the yearly mean tem-

peratures but very slightly or not at all.

3). The pleionian variations of temperature have nothing in common

with the presence or absence of volcanic dust veils.

4). An influence of the sun-spot variation upon the changes of atmos-

pheric temperature is undeniable.

5). In some exceptional cases, at far-distant stations, such close corre-

spondences of the pleionian variations may be observed that one is justi-

fied in admitting that missing crests are due to locally restricted anom-

alies. Of course, in most cases the anomalies are prevalent.

6). A correlation between pleionian and macropleionian temperature

variations and the occurrence and ‘frequency of volcanic eruptions may

be presumed, although, at present, it is difficult to imagine how such a

correlation could be explained.

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‘ a: PRESS OF JUDD & DETWEILER, INC., WASHINGTON, D.C.

ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXVI, pp. 175-214

Editor, EpmunD OtT1s Hovey

STATISTICAL STUDY OF VARIATION IN

- §PIRIFER MUCRONATUS

eee. BY

CHARLES C. Mook

2 NEW YORK ks

PUBLISHED BY THE ACADEMY:

30 JUNE, 1915 |

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum oF Naturat History, 1817-1876)

OFFICERS, 1915

President—GEORGE FREDERICK Kunz, 601 West 110th Street

Vice-Presidents—CHARLES P. BerKEyY, RayMonpD C. OsBuRN; ~

: CHarLes BASKERVILLE, CLARK WISSLER

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[ANNALS N. Y. Acap. Sci., Vol. XXVI, pp. 175-214. 30 June, 1915]

STATISTICAL STUDY OF VARIATION IN SPIRIFER

MUCRONATUS?

By CHarues C. Moox

(Presented in abstract before the Academy, 7 December, 1914)

CONTENTS

“) /Page

ESR BU Ih SS ee Ae eee Neen i re 175

Mutation I.—Spirifer mucronatus mut. alpenense Grabau ms..... PAS 5.5 176

Mutation II.—Spirifer mucronatus mut. multiplicatus Grabau ms.,....... bf 67

Mutation III.—Spirifer mucronatus mut. profundus Grabau ms.......... 179

Mutation IV.—Spirifer mucronatus mut. thedfordense Shimer and Grabau. 182

Mutation V.—Spirifer mucronatus mut. attenuatus Grabau mSs........... 184

OE SNUB ae ga 2 ee Da oe TS ond, os laa veh ws B satin eieupern ahvoia opidwllaie eee webs 186

Paes MCAS ECMOENLS ANG: IWIGICES x. 4.2 occ ceke 6 ope oe ees en sie es we een w elena 190

INTRODUCTION

These studies were made on specimens of five mutations of Spirifer

mucronatus collected by Professor A. W. Grabau and Dr. H. W. Shimer

near Thedford, Ontario, and by Professor Grabau near Alpena and at

other localities in Michigan.

The beds from which these fossils were collected correspond, in whole

or in part, with the Hamilton of New York State. The general suc-

cession of beds in the Michigan region is given below. At the base are

calcareous shales corresponding in a general way with the lower part of

the Hamilton or upper part of the Marcellus of the. New York State

section. ‘These shales contain the mutation of Spirifer mucronatus called

by Grabau alpenense. Above these shales there is a series of alternating

limestones and calcareous shales. ‘The limestones contain the mutation

multiplicatus and the shales contain the mutation alpenense. Above this

series there is another series of calcareous shales containing the mutations

profundus and attenuatus. In the Thedford section, the mutation thed-

fordense Shimer and Grabau occurs in shales corresponding approximately

with those containing profundus and attenuatus in Michigan.

The objects of the studies were to determine, if possible, the relations

of the different mutations to each other, the progress of evolution in the

1 Manuscript received by the Editor 16 December, 1914.

176 ANNALS NEW YORK ACADEMY OF SCIENCES

various lines, and to make a trial of the statistical method im the study

of fossils. :

The curves were made by plotting the percentages of the total number

of individuals having imdices between certain prescribed limits. The

indices were taken as abscisse and the percentages as ordinates. The

higher indices were indicated on the right and the lower on the left.

Progression has been indicated as going from right io left.

The mutation thedfordense was described by Shimer and Grabau in

1902.2 Descriptions of the other mutations are taken to a large extent

from Grabau manuscript.

Muration 1.—Spirtfer mucronatus mut. alpenense GRABAU MS.

This mutation occurs in the lowest beds of the Michigan Hamilton m

a shaly matrix. Ii is of moderate size, with fairly strong growih limes.

The plications are close together and are not very strong. They number

about fifteen on each side of the fold and smus. The shell is sometimes

extended into a pair of mucronate wings. The sinus is broad and shallow

and contains no median plication. The fold is also broad and shallow,

and sometimes contains a median groove in the young stages. There is

never a median groove on the fold of the adult. Three hundred and

seventy-two individuals of this mutation were measured and the follow-

ing is a summary of the resulis. The complete measurements are given

in the tables. ;

Makhnm ‘width. -:. 2.22.5... 0-feeecs beset ae 42mm.

Mininnmm width... ... 2.0230 SS epeseeee le ee eae 15

Maxtmpm length. . 22 2. 62s 2 oe ce te cee ee 28

Minium length. 22 2 22.2265 oe ee ee ee eee oe 9

Maxim shell imicx (adel) 2-222: 224 nS eee eee 2.80

Mintmom shell index (adult): .-.<35-22--2.5-.-2.4s5eeee -95

Average shell index (adult)..-.-....-....-...- (about).. 1.50

Maximum shell index (neanic stage) ..............-.....- 4.39

Minimum shell index (neanic stage)..-................-. 1.29

Average shell index (neanic stage)...........- (about).. 2.60

TapLe I— Number of individuals and percentages of total number of

individuals of various indices

Adult Neanic

yaa Cees ee ;

Number Percentage Percentaze a!

EPR tia hoe e et oc tmeeie 0

1 G1 Se. Fee ere ths Ce eee ee 0

AEs ood Pee eee

A ek es at Ge hres ee

0 TS oe ease cee mee

ae ee

es ea

MOOK, VARIATION IN SPIRIFER MUCRONATUS i Ay

Adult Neanic

Index — AL,

Number Percentage Number Percentage

ey DD 5a) e uote Sere UA oe id 'eitee ane 25 Gut 62 16.6

OO). cis cs eeKeiy = mika eee ae ee a) izes 69 18.5

Lo. ose apacso aus hats © See es 2 a 60 16.2

Sa UPS 0) Mae A eis SER can) ane al 2; 45 12.0

eNO ee N ec ay ac Crm WD Ola OAS siveiel abet ) 0 26 6.9

ee? coe al acoate Wie ac cee mae aia 0 0 21 5.6

PN PAOD: Sol pak ar at day swale or ve @ oats pum Sw 0 0 15 4.0

BME Oe wis sca vaxe wile wie tree ane “ena 8 Gre 0 0 ~ 10

SEM ic). Di eiaee wl bya Gian lee Sis a oe oie 0 0 2 a)

PES IC) ace sic Segcece celal cavaler e's s cos de 0 0 1 2

SW TG tare evens caee o/s aes Aeya Sisk ec 372 99.6 372 99.2

The curve of the adult shell indices of alpenense is fairly smooth. The

shght roughness or irregularity may be attributed to imperfection of

some of the specimens and to the difficulty of deciding, in some cases,

the anterior end of the pedicle valve. The neanic curve is much more

irregular, a condition due, no doubt, to the extreme difficulty of finding

exactly corresponding growth stages in a great number of individuals.

From the size, characters and number of plications, shell indices and

characters of the fold and sinus, alpenense is favorably equipped to be

an ancestor of the mutations profundus and thedfordense.

Moration Il.—Spwifer mucronatus Mut. multiplicatus GRABAU MS.

This mutation occurs in the limestone beds in the Michigan region

above the shales containing alpenense. Shales bearing alpenense some-

_ times alternate with these limestones. It is a large mutation with many

fairly strong flat plications. There are about 17 plications on each side

of the fold and sinus. The fold and sinus are broad and flat, the sinus

bearing a well-developed plication and the fold a median groove. The

growth lines are strong and the young stages are often clearly distin-

guishable. ‘T'wenty-nine individuals of multiplicatus were measured.

RE EET PR VVVEPULS Set cht. 1S ahhicl on aan erie ero eR AN eels PRS Me 50mm.

LTS DUNG COC 7 a1 0 iC AR a Pa 26

MEMENTO OGiC Su. ee mi erick Sk Bee od. eee ee te eee 27

HACAYUIRTOT aT 9 (10 ES eee CS Oc a 15

PSE SHEN INGE (CAG UIE) cols sac gin teeing Sule @ ble hw ence > 2.02

ECON TESST RNS! TV DT 0 C2 a (0210 0 ne 1.46

Averace Shell mdex (adult) occu: c.sasswemweves (about).. 1.80

BaP west SHEL Index CHEAMIC StHHOE) <6... secre eens sewn ns 3.50

lowest shell index (meanic stage)... 05. ccc ce ke ee ek 1.78

Average shell index (neanic stage)............ (about).. 2.65

ANNALS NEW YORK ACADEMY OF SCIENCES

IsuaUad|yD “~nue SNyDUOLONU AafLMdy JO sabnD)S d1UDAU PUD ZINpY UL SadipUL Jays JO suOYIPWOD ArtzDADAWOD BuMoYs saaingQ—T ‘DIyy

SIZF—-/0F

‘00S-94T

SLE—-LE

oS)

00t—9L/

294] —EF/

—~

~™

(

MOOK, VARIATION IN SPIRIFER MUCRONATUS 179

TABLE I1.—Number of individuals and percentages of total number of

individuals of various indices

: Adult Neanic

Index a

Number Percentage Number Percentage

N22) ed L235 | Doe ee mn eee 3 16.3 0 0

eM FN 2a hatenater <yerd came aps <5, sie earene A 31.9 0 0

MNO UN clare! avert. etd aiatoheteinicita G:a la tiers 9 31 2 6.8

ope IE) 315 A ae oS AO en 5 Le .2 3) 17.2

Se ay ie cS a Wenereveiia he elesca a eed e i 3.4 6 2.6

ee EP MN aha schiersre kes 2 cheats a: sh sregau at aye 0 0 10 34.4

Bre By." ona als Bepcuie te Aine, b sete, 206 0 0 2 6.8

Sees ee RE el gw it eyes SIL ein: wien e's o8 0 0 3 10°3

Eameeety ene MU ornate wat ise andy att 0 0 1 3.4

PERO forcible ict, er acral a cs SEhael wach 29 99.8 29 99.5

The adult curve of multiplicatus is narrow and regular. The neanic

curve is broad and very irregular.

ane

ye 34,

/ in : ‘

cane

/ oa :

/ - zoe/ \

as \

/ sche ae

10.37 vs a \ 10.3

“ 9 . <5

| esti ee Pa

(A)- (B)-: s rae

io 2 bor rae

Fic. 2.—Curves showing comparative conditions of shell indices in adult and neanic

Stages of Spirifer mucronatus mut. multiplicatus

Mutation II1.—Spiifer mucronatus Mut. profundus GRABAU Ms.

This mutation occurs with attenuatus in the upper shales in the Mich-

igan region. It is small with extremities not at all or only slightly

mucronate. The median sinus is deep and angular. It extends from

the beak to the anterior end, then beyond into the brachial valve. The

bottom of the sinus is subangular. The fold on the brachial valve is

shallow and has no median depression. The plications are round and

close together. There are 8 to 12 on each side of the fold or sinus,

180 ANNALS NEW YORK ACADEMY OF SCIENCES

rarely less. The growth lines are strongly turned back over the fold

and sinus.

Mehr a yh Elsner ree ene ee te aa 35mm.

Miniimiim+ awad tins tec exesee ese re ene ee tee ee eee eee Sieue 15

Maximum Wemerthics eke ers send ees a cai eae retene 21

Minimum feneth:3 3 che ao ies cee ae eee eee 9

Highest: Shell index -(adult) fancies oe nee one eee eee Pape

bowest shellindéxs (duit) 325. eee ee eee eee 97

Average shell index (adult)......... eee eee (about)... 1.65

Hichest shell index, (neanie stage): .-2..cen = are eee ae 4.37

iowest shell index (meamic staze).-2.. 4-400 cee eee 1.35

Average Shell index (neanic stage)............ (about)... 2.50

The adult curve is fairly short and regular. It corresponds closely

with the adult curve of alpenense, though the average index is a little

lower. The neanic curve is very broad and irregular. It corresponds

rather closely with the neanic curve of alpenense. ‘The average neanic

index, however, is a little lower. Two hundred and seventy-six indi-

viduals were measured.

TABLE III.—Number of individuals and percentages of total number of

individuals of various indices

Adult Neanic

Index (ea ee —

Number Percentage Number Percentage

SOO. aie S sapere nes Seoeeem enters 2 7 0 0

} OASIS ZB, recite tk ere ec a ager 36 13.0 0 0

1 ZO Ad DO ee ah ee eee are ee eee eas 112 40.5 2 Bart

he aera as Say seek eae eae: 75 Zt Al 6 pa |

LA O22 B00 se sidtovaee ie ere Cerne aici 40 14.4 37 13.4

Pr ONE De siar opeeta oa ea eee ste eee teers 6 rea | 72 26.0

Di PAGE OO ors Ek SRE te eating Sete 3 1.0 52 18.8

Dis By NEDA < velianeiscusvsanon Ovepeaseaae se eeunercgane ie 2 at, 60 21.5

2 ABS LOO! s UPS, GeO Reeves arene eases 0 0 31 11.2

3 OISS 22) ceo oe ee ee eae 0 ) 8 2.8

DOS OO eeetesanien svete aera Ss euaegeate eee 0 0 = 1.4

So Chl es ALO ai austa Cle rord esters ate es tate eee see 0 0 0 0

BOAO aoe nein eee oe ee eee 0 0 2 eit

A ON AAD hig Ste ilo whe Serene Cee ues 0 0 il. 3D

42GB SOF ogc alk aie whe oA 0 0 1 .o0

Morals. Wurcs: canieeeas is ead oe 276 99.5 276 99.3

This mutation is evidently an offshoot from the alpenense stem. The

line of development has been more in the nature of a deepening of the

sinus and strengthening of the growth lines rather than in reducing the

shell index. The shell index is a little lower than that of alpenense in

181

MOOK, VARIATION IN SPIRIFER MUCRONATUS

snpunfoud “Qmik SsnaDWOIONU MaJIadgy JO SaHN]8 OUDIW PUD WINpYD We sadrrpU Yays Jo suo IpUOD dAYDADdWMOod OWMOYs Sari O—"e “DIT

+ w w cA w is) bi w re) =< a a 2x4

be g R 8 g x a ; x a & s

t ' 1 | 1 t 1 ‘ J ' |

- - - w w w +) LS) b b = =~ —

haere : : 5 : : 5 : | : : 3 ee

2 a i: o- a ai hg ;

“bt

182 ANNALS NEW YORK ACADEMY OF SCIENCES

the adult stage, but only a mere trifle lower in the neanic stage. This in

itself is an indication of progression toward a low indexed condition.

Mouration 1V.—Spirifer mucronatus Mut. thedfordense SHIMER AND

GRABAU

The mutation thedfordense occurs in the upper Hamilton shales im the

region about Thedford, Ontario.

This mutation is the most advanced of the five mutations studied. It

is fairly good-sized and very stout. There are 12 to 15 strong sharp

plications on each side of the fold and smus. The sinus is deep and

subangular in the adult and occasionally has a faint elevation in the

very young stages. The fold is somewhat elevated and rounded in the

adult, and often shows a faint median depression in the young stages.

The growth lines are very prominent, and are sharply deflected over the

fold, sinus, and plications. Two hundred and thirty-one individuals

Were measured.

Maxinigm: wiGth. ...'3 <25< 2 si. oo eee eh ee ee eee 45mm

Maximum Tengtlt oo 2253 pos See eee ae ee eee 35D

Minimum feneth) . 22; 22 ee ween ee eee eee eee eee 15

Highest shell index: (adult) co2.. = ae oe ee ca ene eee 1.94

Lowest shell“index’ (adult). 2. fe2n se ae oe cl eee 89

Average shell index ¢adait) . veo 26 54-0 eee te (about).. 1.30

Highest shell index (neanic stage)...........-.........-- aoe

Lowest shell index. (meanic stage)... 06.2... . .s'-. « 22 eee 1.24

Average shell index (neanic stage)............ (about).. 2.00

Taste 1V.— Number of individuals and percentages of total number of

individuals of various indices

Adult Neanic

Index (pee

Number Percentage Number Percentag

= OO oo ob. Sis oe ee eres 24 10.3 0 0

AE 2S fw oe We ee ee ee 107 46.3 2 Cae:

2 2D BD). 3 ao oaths bine eee ees 80 34.6 17 tock

Be TE oii See oe ee ee eee 15 6.4 54 23.3

PEGS OD oh seed Be eee ee 5 2.1 63 26.8

DE os Seiden = on eee ee 0 0 40 17.3

2 BD nt & ues ee eae eee 0 0 24 10.3

RD) SiS hae Dine Sore ew See ee ee 0 0 18 t.8

2 FGA so ok es Jae eee ee 0 0 10 4.3

SS OES 3S 8 kd etek tes eee ee 0 0 -: 4

5 ap, ae ae | eee ee Sees We LPT S 0 0 1 .4

Cet ee Sy ee eer ee a 0 0 1 4

231

bac]

DN)

—

pot

OSD

SPIRIFER MUCRONATUS

MOOK, VARIATION IN

asuapso{payy “gnu snqouowonm safindgy fo sabn7s ovunau pun 4yjnpy UW saavpuy qjays fo suo1Ipued daYnundwmo0d HBuMNoYys SAA~AND—F “DIT

OSE-FTE

*Hesee -/08

'00E—-ILZT

;

S2r-loet

000-941

0.$[-Fe/

-00/-

184 ANNALS NEW YORK ACADEMY OF SCIENCES

The adult curve is very narrow considering the large number of

individuals measured. It is regular in form. The neanic curve is fairly

regular and is very wide. It would correspond fairly well with a flat-

tened-out curve of the adult.

Thedfordense was evidently derived from alpenense along with pro-

jundus. It differs from profundus in having vestiges of a plication in

the sinus and groove in the fold, in being more robust, and in having a

lower shell index. Profundus has progressed further in the deepening

of the fold and sinus, and thedfordense further in reduction of the shell

index.

Mutation V.—Spirifer mucronatus MUT. attenuatus GRABAU MS.

Attenuatus occurs in the upper shales in the Michigan region along

with profundus. The shell is very wide and short, giving a high shell

index. The sinus of the pedicle valve has a distinct median plication

which extends nearly to the beak. The fold of the brachial valve is flat

and level with the plications on either side or slightly elevated above

them. It has a pronounced median groove which extends nearly to the

beak. The growth lines are moderately strong. There are 15 to 18

strong plications on each side of the fold and sinus.

Maximam-wiith.: 5 sé. ..0. tee as ees hee oe eee eee eee 47mm.

Minimo width: o s/s 23 SEC 2 ation See ee ee eee eee 19

Masimum Jenpiiie <9. 2 ¢eet Soe eee Ce ee oe eee eee 17.5

Minimum Jeneih: 2 pods. eee See ae eee bee eee 8

Highest shell index (adult) 22:02. S.. c.ce2s Jes sector 3.75

Lowest .shell index \(adett) 2202.75. ee ees ou. ee eee ay)

Averace shell index -(adult) a2 02. eae eee (about)... 2.25

Highest shell index ‘(neanie stage)... {2622 25.22. sec Seca 4.70

Lowest shell index (neanic Stage). ..252-...2.s-c..eeeee yA

Average shell index (meanic stage)............ (about)... 3.40

TABLE V.—Number of individuals and percentages of total number of

individuals of various indices

Adult Neanic

Index ed

Number Percentage Number Percentage

1 Pe ce od Sk so ee eee 2 2.0 0 0

6 ed ni en cele cee eee 7 9.4 0 0

De oe Recs ee aie eneiie me 16 21.6 0 0

2 2 Rs ak Swe we ok eh oe eekoae 1s 24.3 1 1.3

2. ne ome See ow ae eee oe 14 18.9 6 8.1

2.2 TE: code oe ates eee teens 10 13.5 7 9.4

2 0D. ie oct as vs kee een eee - 5.4 13 17.6

oo a

do ds

ce)

bape te)

“1

Ht ©

veo)

185

MOOK, VARI y

ATION IN SPIRIFER MUCRONATUS

SNYDNUAIAD *

1p “Qn Sn.DWO

wonw dafiud

wdg jo sabn

18 OVWwDAW |

1 pun .)npD

UW SadUpuUr 1

Q yur rays jo s

WOVUApUoOd a:

yi “UzwIvawUo0d b

1 9 Buaror

1 {S SOLMMQ—

VAN Q—G “OTH

a ra

C+ ae eee

: . E ‘ 3 by q

: s 8 : N

: 1

7 i v Fr x bs ®

3 x 5 : i :

“yr : 4 J Ne & Ve 1 < x a e

a ——— e s x S l ri q &

AA Ree: q Non CDAD RE eee ° u S : a : :

ae 7. Lv 3 :

9 Ny Z .

Ligh |

A 3

ae +s. ae

“ : 3

a 7 c* 2

79 3

i) aia |

iD at hey aan eo V8 =<

ta po Vel ara ue ,

ate ale r a le sets .

nae Fae ioe ae AS

bet

eras cs

cay ‘

aS ‘“ i

age coe

186 ANNALS NEW YORK ACADEMY OF SCIENCES

Adult Neanic

Index ee

Number Percentage Number Percentage

3 .b1-S. se coe oe ee eS ee 1 1.8 10 13.5

31642 00 25 eee ee oe Eee 0 0 ) 6.7

4 O14 2S ih ees He ee SE 0 0 2 raf

B24 BD Ss ae Se Eee See 0 0 » 6.7

4c Dl (De oee SSEe Soeeeeeee 0 0 2 2:7

otal on ces ote eee ee T4 99.8 74 99.7

The adult curve of attenuatus is broad and rather irregular. The

neanic curve is very broad and irregular.

This mutation is in many respects the most primitive of the five muta-

tions studied, although it occurs in the highest of the three horizons. It

is nearest to multiplicatus in form and general characters though multi-

plicatus has a lower shell index.

The derivation of attenuatus is uncertain, though it is not far removed

from multiplicatus. Several possibilities may be considered. Attenuatus

may be derived from some form of muj/tiplicatus whose index is much

higher than any of those measured. Jt may have come from a multi-

plicatus stock having an index like the highest of those measured and

then degenerated into a short and broad form. It may have sprung

from a common ancestor with multiplicatus in a lower horizon and re-

mained primitive while mwultiplicatus progressed toward a lower index.

The third possibility is by far the most probable. The adult curve of

attenuatus corresponds roughly with the neanic curve of multiplicatus.

CoNCLUSION

Two distinct lines of development have been observed in Spirifer

mucronatus. In one of these lines development has advanced to a con-

siderable distance from the primitive Spirifer mucronatus condition.

This line includes alpenense, profundus, and thedfordense. The other

line has progressed much more slowly. This line contains multiplicatus

and attenuatus. In the first line thedfordense and profundus have prob-

ably been derived directly from al/penense. In the second line attenuatus

was probably not derived directly from multiplicatus, but from a form

closely resembling mu/tiplicatus, but more primitive. Multiplicatus and

alpenense were probably derived from some primitive form of Spirtfer

mucronatus belonging to a lower horizon than any of the five mutations

studied. The approximate relations of the five mutations to each other

are shown graphically in the following diagram.

MOOK, VARIATION IN SPIRIFER MUCRONATUS 187

attenuatus profundus thedfordense

multiplicatus

alpenense

Primitive form of Spirifer mucronatus

Fic. 6—Diagram showing the relationships of the five mutations of Spirifer mucronatus

The tendency in development has been to reduce the shell index, to

reduce the number of plications,-to lose the groove and plication on fold

and sinus, to strengthen the growth lines and individual plications, to

deepen the sinus, and to a certain extent to reduce the actual width of

the shell.

ANNALS NEW YORK ACADEMY OF SCIENCES

188

Snynoydyinm ,,

9 99

PSE) O10 ase 8 0) a snpunjoud “yn

snpouosonm safimidgy Jo suoyngnu auf fo sabnys ynpp ay. Jo

w ics} & A :

4 g = iS 4 R

[ ! \

cy) Gy A)

3 : iy 8 5 3

HON a

5 Cael}

5] AiNiainietn ar raiione Gen AR

Le *

”

—- — — — — asuauadyo ,,

99 99

‘xopul yjupe = ———_———____ 9suapuofpay, “ynm

saoipur ays JO suonipuod —daynsndwmo0d HmMnoys saavgo— ),

00% - 9L/

97 (-/S/

S7/-10/

ool

‘DIA

189

SPIRIFER MUCRONATUS

MOOK, VARIATION IN

” ” RA RA ARARARA 8NYDNUAIIO ”

ij a) 0: eee ee — snqvoydyinw ,, vs x =o asuauadjo ,,

‘SOOUP OP MUO) cease naeeseumeeces snpunfowd "yn ‘xopuy syuvou =——_——____—___ asuap.sofpay? *ynu1

snpnuowonu Jalrudgy fo suoynenw auf lo sabnys aunau UW saoipur ays fo suo1Nypuod aaynsndmos HBumoys saainp—s “DI

S &

cr el

“00 f-9AT

YG ded AY

‘002 -9L|

StE-—10e

‘Stl -}9/

190

eeeee

see ee

eceeccece

eeceee

or)

Co Co CO

ea)

ede

oeeee

eeeee

ceaeee

«eeee

oeeee

©“eeee

“eeee

ANNALS NEW YORK ACADEMY OF SCIENCES

‘TABLES OF MEASUREMENTS AND INDICES

Npirifer mucronatus mut. thedfordense

Adult Stage Neanic Stage

(Es si =e aN (Ge

Width Height Index Index Width Height Index Index

(reverse) (reverse)

Simm, 929mm 72935 iL. O06 Simm, 23mme Sc: 1.34

32° 29.5 921 1.08 32 Z0es .640 1.56

Bye ie .843 LS 32 AT, Sea 1.88

38.9 27 Sabi 1.42 38.5 20.5 02 TSG

30 3 1.000 1.00 3 24 . 800 1.25

29 ZTE) 948 1.05 29 eas O72 1.48

31 26.5 bd 1.16 31 19 612 1.63

3+ 28 .823 te, 2. 54 23.9 .691 1.44

om Bee 1.015 .98 32 18 . 962 WAAAY

2 2B Bs ail! lial QT die .629 1.58

54 24 TOD 1.41 54 a9 OOS 1.78

39 530.5 871 1.14 39 19 542 1.84

39 28 . S00 1 WBE) 35 Ay, ey .500 2.00

38 24-5 . 065 1.76 38 16.5 434 2.30

5- Pit . T94 ha2b 34 Ze 647 1.54

30 26 S66 JERR) 30 20 . 666 1.50

Ste (e539 723 8's) 38 18 473 pipes 1

32 28 S75 1.14 on 20 .625 1.60

29 De 793 126 29 1935 “G2 1.48

33 2ORS 893 alee slat 33 48.5 .560 1.78

29 24 827 1.20 29 IPT Ges) .603 1.65

30) 25 .t14 1.40 oD 18 514 1.94

36 3 S61 1.16 36 aaah .583 ge (a.

28 24.5 S75 1.14 28 17 . 607 1.64

ita) 26 742 1.34 35 Pepe .628 1.59

32 23-5 T3834 Weals Be a L7G m5 )3) | 1.88

Pat 2 851 2 ae of 19 . 7038 1.42

29 aH | 931 1507 29 Di . 124 1.38

40.5 28 .691 1.44 40.5 ZO KES 506 1.92

Beas, 3 . 923 1.08 ao Je a Wega: 038 1.85

28 19 678 1.47 28 15 585 1.86

a1.5 Zed 904 1.10 3) Na) 18 ay ial ies

38 50 7TS9 1.26 38 23 . 605 1.65

26 20 740 tao AT eas 14 .518 1.92

35 23 657 1. 52 50 18 514 1.94

31 20 645 dea ak 14 451 at I

Ot se 26 693 1.44 S120 20 N30 1.87

40 32 . 800 120 40 2a 575 Lis

mk A) * 806 1.24 ou VE 548 1.82

a2 y) 906 1.10 32 19 .993 1.68

Number

Peeves

aS eS es es S|

He wwe

eeceese

eeeee

MOOK, VARIATION IN SPIRIFER MUCRONATUS

Width

Spirifer mucronatus mut. thedfordense—Continued

Adult Stage

—

Gn, SE Ee aE

Index Index

Height

(reverse)

26mm. _ .896 pa Ba:

3 . 967 1.03

29 1.054 .94

25 .781 1.28

26 . 896 tM

25 hee 1.32

24 . 750 1.33

25 . 833 ¥.20

22 745 1.34

26 702 1.42

ra | .870 1.14

24.5 . 662 1.51

24 . 800 1.25

25 tSk 1.28

21 750 1.33

23.5 734 1.36

26 787 1.26

23 . 851 eke

26.5 . 883 iis

26 . 962 1.03

23 .676 1.47

25 .625 1.60

24 Ae of 1.37

30 . 769 1.30

24.5 . 844 1.18

21.5 .53T7 1.86

29 . 125 £-3¢

27 .710 1.40

21 . 750 1.33

26 . 742 1.34

25 .641 1.56

29.5 . 867 1.15

hs ae 1.23

25.5 . 980 5 AT

24 .8138 1.22

25 . 862 1.16

25.5 . 879 1.13

25 . 806 1.24

16 ool 1.81

15 . 930 1.86

21 . 646 1.54

25 . 806 1.24 °

20 -588 1.70

Height Index

(reverse)

20.5mm..706

PA! 677

20 Be fs

18 . 962

17 .586

Ve 015

19 .593

18 . 600

14 474

18 .432

20 . 645

15.5 .418

19 . 633

15.5 .484

16 .OT1

18 .562

19 075

15 .055

17 .566

16 .592

18 .529

19 -475

7 515

20 512

18.5 .638

LG25 437

24 . 600

20 my AS

a rg . 607

LFS .500

18.5 474

21 617

20 . 625

15 .576

13.5 457

SES5 . 603

15.5 .596

i .548

12 .413

oF -392

17.5 .538

18.5. .G29

15 441

Neanic Stage

191

Index

ce oN Oe GO

el el 0 oll oO OSS

“I-91 WwW Ap

Soa a al

192 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. thedfordense—Continued

Adult Stage Neanic Stage

— aay SS

Number Width Height Index Index Width Height Index Index

(reverse) (reverse)

‘oie Iperenieee 33.5mm.26.5mm. .791 1.26 33.5mm.20mm, .597 1.67

SO eS 28 PALA) acre 1.30 28 17 . 607 1.64

GO keer 30 30 1.000 1.00 30) 19 . 633 1 52

os ae 34 25 735 1.36 34 18 .529 1.88

Se 39 27 . 692 1.44 39 19 .482 2.05

ORs cris 34 26 . 764 ool 34 19 .5d8 a eT fe:

oe age 34 25 pasa) 1.36 34 19 .558 148

Oy Le ote 34 23 .676 1.47 34 14 .411 2.42

SG ieee 43 29 . 674 1.48 45 19 441 2.26

a earerge ol 30 . 967 1.03 31 18.5 .596 1567

Seo ees ao 24 .685 1.45 35 aS .500 2.00

OO ere isie DAE 26.5 .981 1.01 2 19.5 tae 1.38

HOO S. se5 39.5 Bye) . 822 gyal | ae 19 O84 ae:

101-175 inclusive — Spirifer mucronatus mut. alpenense.

PAS Mae 33.5 32 952 1.04 33.5 20 597 1.675

P| V2 eae 20 Dianed . 944 1.05 2a 14 .518 1.92

DM £5 Fee ae 33 . 968 1.03 32 19 .593 1.68

AG fe ae 26 28 1.076 .928 26 18 . 692 1.44

ZA We Se? 30.5 28 .918 1.08 30.5 14 .459 2. Mik

BES S08! 33.5 32:5 .970 1.03 33.5 20 .D9T 1.675

51 eee 26 26.5 1.019 .981 26 14.5 Dot 1.49

yd 1 eA 35 33 .942 1.06 35 22.5 .642 1.55

pit 1 RN Laas Zt Zit 1.000 1.000 27 18.5 .685 1.45

DEAS borows Si 29 - 985 1.06 31 20 . 645 1.55

PaaS Brae 26 2 . 961 1.04 26 17.5 . 673 1.48

De er sats 28 26 .928 1.07 28 2st) . 803 1.24

Fy hove eis 29 26 . 896 Kolt 29 20 . 689 1.45

PAG gees By 31 . 968 1.03 oe 22 .689 1.45

227-445 inclusive — Spirifer mucronatus mut. alpenense.

446..... 39.5 26.5 .670 1.49 39.5 iy G5) .4438 2.25

447..... 3 pape . 666 g Meta f axed 14.5 .439 pee F

SAS es. 35 ok .885 q oe 35> 17 .485 2.05

449..... 36 291.5) . 708 1.41 36 20 .555 1.80

ABO te sss 36 22 .625 1.60 36 16.5 .458 2.18

7 15 Ore esis qe ye ee echt heme «tates esta

4B nos a 33 23.5 th 1.40 33 16 484 2.06

ADS ania a's 37 28:0 770 aL 29 37 18.5 .500 2.00

454..... 30 24 . 800 L.25 30 17.5 -583 oe fs

ADD os os. 34 27 . 194 25 34 22, 647 é Aa:

co Oe 36 AL a . 763 1.30 36 20 556 1.80

AML 5.0 65 32 221.9 . 703 1.42 32 14.5 .453 2.20

BS ot anccs 35 27 errs 129 35 19 .542 1.84

4D oc0 5 31 25 . 806 1.24 31 18.5 .596 1.67

AGO. PS ci 45 30 . 666 1.50 45 21 . 466 2.14

MOOK, VARIATION IN SPIRIFER MUCRONATUS 193

Spirifer mucronatus mut. thedfordense—Continued

Adult Stage Neanie Stage

| aa -A~~ a FSA, =e <a ies. - =, Dae

Number Width Height Index Index Width Height Index Index

(reverse) (reverse)

AGI oc.) s6mm. 28mm. .177 1.28 36mm. 2ilmm._ .583 Bat

AGA... s 32 29 . 906 £10 32 19.5 . 609 1.64

AGS. 30 » 33 28 . 848 iL Galrg 33 18.5 . 560 1.75

464..... 35 26 742 1.34 35 Al .486 2.05

465..... 30 Peas ATO) 133 30 TGS .550 Loa.

2 Caer 33 2.0 eee 1.40 33 18 545 Uae

GT. tes: 36 23.5 . 652 dos 36 16 444 Zao

PRs a, « 30 24 . 800 b25 30 16 Jao L815

459-605 inclusive = Spirifer mucronatus mut. alpenense.

G0G6..... 32 By 1.000 1.00 32 18 .562 g wr are

TG ss ss Bol at 1.000 1.00 Sil AG 516 ese

GOS. 2... 23 aa 1.086 .92 23 eh. 478 2.09

GO9..... 28.5 25 Oe 1.14 28.5 At 526 1.90

ClO oe. eee ara ts Resets sa etal Wee Pita bf

C13 30 28 . 800 As: oo 13 ew: 2.69

Gis... 33 26 ae (316 26 aa 15 .454 2 20

GHD 02. 36 31 . 861 ipSale 36 20.5 .569 1.75

ol eee oe esses Me Beet tae 3 pavers fe eae Bes

OMG ioe... 36 30 .833 2220 36 14 388 2.56

GBs oe. 3 aie 5 20.0 . 746 1.34 os ATS BETS: 1.80

Got. ba, 's2 hess 3 Ser waits feces Sipev's sept oceke

2 | ee 38 eee .828 120 38 18 .473 2.11

Bele Pare Fees snatine Stisate aes gs ae jcgaee

RU ees a 20 29 1.074 .931 2 16.5 .611 1.63

Goan 5 «6 Dee 20.5 S7alae 1.14 23.5 12 510 1.95

6244 0... Be 25 781 ies 32 16 .500 2.00

Goan... 31. 28 - 903 10 onl: 18.5 .596 1.67

GAG... ., eieaers neh teieae ake hu By ys Shas seeate te te

27 ra 34 Ses .926 1 OG 34 1.5 .450 2.19

Geaicin a cs 31 26 . 838 pe 31 12 O06 2.58

i ae Bes sae exis ae ee sifade Soke es te aa

GeO... s 30 30.5 1.016 . 983 30 18.5 .616 1.62

ieee os 28 20 .714 1.40 28 a (55 .410 2.43

2 32 29.5 921 1.08 32 18 . 562 gar ar

Deion... 30 20 . 666 1.50 30 ils .433 2.30

Ga4..5.. PA | LT, .629 1.58 20 15 426 ives

io 30 18 . 600 1.66 30 10 333 3.00

Ga... Zhe 24 802 1.14 2h tb 490 2.05

Gals... ot.5 26 .693 1.44 St .5 a, .4538 2.20

C2): ne 2.0 ah . 823 a 8 VAT ES 10 392 Ds ees

oo. eae oh 21 . 743 3 ee ot 1% 459 2.17

640..... 27 28 LOST. . 964 At 16 . 592 1.68

2 ee rs oe .888 pe Biss 36 18.5 RTs fe 1.94

G2. .... 2k 26 . 962 1. 08 2 10 .370 2.70

00 at 28.5 1.055 947 27 16 .592 1.68

194 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. thedfordense—Continued

Adult Stage Neanic Stage

ee eg

(reverse) (reverse)

644..... 3 . 28mm. .903 1.10 3simm. 15.5mm..500

Gt sees 27 23.5 .870 1.14 2 10.5 .388

G46. 5: aa 26 . 962 1.03 _ 2a 13 .481

G47 2 Sse: oe anes on = wees Pie oats

G455 >>. 29 17 986 1.70 29 10 344

Gik 2 oe 32 35) 1.093 914 32 15 468

G2. oS 29.5 26:5 S9S teat 29.5 14.5 491

5 eee ae 354 1.096 wi! i Bf 31 Hy .532

le eee 38 31 $15 122 38 13 342

Geet cs 31 25 S06 1.24 31 255 403

Gabi 25 ys 1.040 .961 25 11 440

GaSe 2.x. 30 31.5 1.050 952 30 16 503

it) eee 24 20 33 1.20 24 9 379

1 sae rt | Zico ss 981 27 12 ait

te. 37 19 514 1.94 37 11 297

GS...) = 27 S70 1.14 3 12 387

664..... a

Giese cs 30.5 20.5 672 1.48 30.5 13.5 442

666..... mpegs of? ve F

TCT ire 30 29 . 966 1.03 30 15.5 .516

Pee 24.5 A oh). ASSES .89 24.5 14.5 . 604

7 TS See 27.5 27 .981 1.01 21.5 16 581

Gre. = 1G: rf 93 1.14 = a 16.5 523

ye im hae 30 2L.5 716 1.39 30 15.5 516

Gia Ss S41 yf (ee SST 1212 si. 14.5 467

Eee 40.5 27 666 1 50 40.5 16 395

ifs ete 31 24.5 790 1.26 31 17 548

1s eegeaione 3 24 .610 1.62 Sy 16.5 423

Giles: 35.5 3 .§45 1.18 35.5 17.5 .494

yt Rae oe 3.5 27 .620 1.61 43.5 17.5 .402

ire. ox cae 26.5 736 1.20 32 EGS 343

yee 31 23 .D 754 1-31 31 1 .419

ic) ee 39 28.5 i3 cE 39 16.5 428

4 eee 37 25.5 .689 1.45 37 14 378

6S2 40 28.5 712 1.40 40 13.5 437

Git 2% 34.5 25 724 1.38 34.5 14 405

684..... 34.5 ry 197 1.25 34.5 15.5 .449

27 ee ae 37 29 i83 i 37 16.5 445

ites =: ae ot 25.5 689 1.45 37 14 378

oy aa 32 32 1.000 1.00 32 18.5 578

12 ae 38 26.5 .697 1.48 38 13.5 .355

2 ee 31 28.5 919 1.08 ol 17.5 - Dae

ci | iti 54 23.5 691 1.44 34 14 411

MOOK, VARIATION IN SPIRIFER MUCRONATUS 195

Spirifer mucronatus mut. thedfordense—Continued

pdult Stage Neanic Stage

Number Width Height Index Index Width Height Index Index

(reverse) (reverse)

Coe eee 36mm. 2imm. .7T77 a Ae: 36mm. 15.5mm..43 2.32

GO2e 6 29 26 . 895 cee 1 29 14.5 500 2.00

BOG saws Sa <0 Diy .T61 io ao 12.5 “ote 2.68

OS See 28 pee | . 964 4303 28 14 .500 2.00

CS ae 35 29.5 .842 #48 35 af .314 SAT

BGS. isis < 31 22, . 709 1.40 31 i 5 skO 2.69

Co ee 37 22.5 .608 1.64 37 t225 .337 2.96

ols eae hes Seis ee Sey ate te

tS ieee 39 26.5 .679 1.47 39 13 .330 3.00

0 er 30 30.5 | 7 1.016 . 983 30 15 .500 2.00

oe 30 23 . 766 1.30 30 13.5 .450 Zee

le 42 34 . 809 2S 42 15 soot 2.80

Rates's s ee BS os ote ee A eee aenke wet Seah

Wate... 36 20 . 805 1.24 36 13.5 mY (3) 2.66

(A ee 31 aL.5 1.016 . 984 31 14.5 . 4038 2.43

7.0 31 29 .935 1.06 31 155 .435 2.29

7) ae ee sicher =e ses Bes: eae d

DIOS 6 ose = 35 23.5 Boy! 1.48 35 ib 428 2.0

“(tS 34.5 28 .811 1.23 34.5 18 O21 197

iC) an 31.5 26.5 . 841 1.18 31.5 15.5 .492 203

ti 30 24 . 800 Rees 30 ee .416 2.40

oe 36 so.5 1.0138 . 986 36 18 .500 2.00

WAS es 3 ss 36 29.5 .819 122 36 16 .444 2.25

ee... 33.5 29.5 . 880 1.38 33.5 15 447 2 De

Aas. Ss 29 26.5 913 1.09 29 15.5 536 1.87

Gos: . 33 29 878 iets 33 14 424 2 3

‘i a 36 30 833 1.20 36 13 361 2.76

“ae 40 ry es) 687 1.46 40 15.5 387 2.58

Ge. os 41 29 (07 1.41 41 TL5 .280 3.56

(a 38 29 763 fi 23 38 14 .368 2; Th

i) Sa 41 29 .3 714 38 41 16 .390 2.56

ees. Sor bots Sahreln ehectvs tyne ogee

Mea ss 29 25 862 1.16 29 13 .448 2235

Spirifer mucronatus mut. alpenense

_. ee 38 22 918 £272 38 14 oOr pApsy fi |

: 34.5 20.5 578 72 34.5 125 . 362 2.76

ee... 35.5 19.5 . 549 1.82 35.5 13 . 366 28

mete. Ss 34 A 53 easy 34 12 -nehen 283

Re ss. 32 pa a] 640 156 32, 14 437 2.28

2. 29 17 586 5 ar 29 12.5 .431 2232

ees... 38 1-5 D138 1.94 38 12.5 .328 3.04

Lae 27 16 592 LS AT 11 . 4038 2.45

i ae 32 23 718 1.39 32 15 .468 24428

196

Number

3 = (sei te

“eevee

*| = 6) os

"ee ee

CPeevoevee

“ee ee

= se ee

w a terel es

ae a ew

Width

Adult Stage

Height Index Index

(reverse)

22MM, iso 1.36

20.5 . 500 2.00

did oe .442 2°26

20.5 . 640 1.56

17, . 680 1.47

18 .620 1.61

19 642 f 263

a eS ool d voy s

18.5 .486 2.05

Be . 647 1.54

22 .68T 1.45

24 . 600 1.66

i) .500* 2.00

22 .o94 1.68

20 .506 LIF

Zs . 833 1720

23 ese 1.34

21 617 1.65

23 657 1.52

18 . 692 1.44

ps 3) Bu, 514 1230

23 - 93 1.26

22 785 127:

13 . 464 2.15

23 . 884 TAS

i | . 700 1.42

21 677 1.47

195 973 1.74

25 961 -1.04

23 . 605 1.65

23 . 766 1.30

20 . 689 1.45

25 .TS1 1.28

22 .814 ti 22

21 . 636 1.57

16.5 932 1.87

22.5 . 803 1.24

26 .812 1.238

20 . 666 1.50

13.5 540 1.85

20 . 83: 1.20

a . 124 1.38

12 .415 2 deh

‘Width

30mm.

ANNALS NEW YORK ACADEMY OF SCIENCES

Npirifer mucronatus mut. alpenense—Continued

Neanic Stage

Height

Index

(reverse)

15.5mm..516

—

a i

OT OT OF OT OT

. 292

. 288

.468

. 460

344

387

084

ww.

Ww:

OR Ne Od Se)

NOWwWnWwnwnnwr Ww wwnpew

Www ble bo

Nw eH &

wNomwWM Wh WY Ww bw

Wwe bd be

MOOK, VARIATION IN SPIRIFER MUCRONATUS 197

Spirifer mucronatus mut. alpenense—Continued

Adult Stage Neanic Stage

Width Hane Index Index Width Height . Index Index

(reverse) (reverse)

30mm. 22mm. .733 1.36 30mm. 14.5mm..483 2.06

54 3 -382 2i Gl 34 S .235 ote

29 15 ea G 1.938 P49) el? 318 2.63

26 14.5 ont 1.79 26 10.5 p11 2.47

oe 16.5 912 1.938 32 12 oto 2.66

34 18 -029 1.88 34 12 .352 2.83

30.5 23 .613 1.62 37.5 13 . 346 2.88

34 18 .529 1.88 34 10.5 .388 3.23

26 15 016 1.73 26 8.5 .326 3.05

29.5 17 576 1.73 29.5 10.5 3 193) 2.80

3 19.5 629 1.58 31 IZ .5 .403 2.48

on 22 687 1.45 32 13 .406 2.46

28 26 928 AO7 28 13.5 482 2.07

26 18.5 711 1.40 26 10.5 . 403 2.48

32 27 . 845 1.18 32 16 .500 2.00

26 23.5 . 903 1.10 26 AT . 653 1.52

15 9 . 600 1.66 15 ic .466 2.14

29 17 .586 1.70 29 cia .379 2.63

30 18 600 1.66 30 ala! . 366 2.2

ok 20 645 1.55 31 15.5 .500 2.00

36 18.5 5138 1.94 36 12.5 .346 2.08

29 18 620 1.61 29 11.5 .396 2.52

37.5 22 O86 Bef 37.5 11.5 .306 3.26

23 24 1.048 -96 23 14.5 . 630 1.58

yy 20 909 1.10 22 B35 .522 1.91

39 23 589 1.69 39 16 410 2.43

39.5 16 405 2.46 39.5 11-5 291 3.43

29 22, 758 gtr 6 20 16 551 gE |

32 19.5 609 1.64 32 13 406 2.46

38 15.5 407 2.45 38 ae 289 3.45

30 3 .433 2.30 30 9.5 316 3.15

or 18.5 596 1.67 oo 14 451 2

36 20.5 569 1.75 36 138 361 2.76

28 19 678 1.47 28 13 474 2.15

36 18.5 513 1.94 28 on5 410 2.438

32 21 656 1.52 32 14 437 2.28

28 15 .5385 1.86 28 12 .428 2.38

28 22 785 1.22 28 9 321 3.11

ah)

fan)

eo)

ey |

bom

bak

yea

fan)

feed

bed

=r)

bem

fom)

ses ee

= P'S ins

"eee

"see

*= es @es

Adult Stage

Width

Height Index Index

(reverse)

16mm. _ .533 1.87

28.5 .950 1.05

20.5 .500 2.00

20 645 1.55

19.5 .513 1.94

17 43 2.05

21 538 1.85

24 750 1.33

21 717 1235

20 588 1.70

21 .750 1.38

28 .875 1.14

22.5 sis i

19.5 .750 Be=

Ws 653 1.52

17.5 .480 2.17

23 621 1.60

19.5 .590 1.69

24 827 1.20

20.5 .525 130

22 . 709 1.40

raysss) 1.80

19 .452 2.21

25 .851 i Bea Wg

18 . 600 1.66

21 . 700 1.42

20.5 .625 1.75

19 .465 2.15

Width

30mm.

2 dS Ww oo

ip)

=r)

“I ND 0

ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. alpenense—Continued >

Neanic Stage

Height

13mm.

14.

or Ot

19) |

Or Ot

Index

(reverse)

.433

.516

.990

. 467

.451

.328

271

294

NW po

bo = Oo

me WwW OD

9 9

9 19

09 NY ty

BSE

hm OD OO be

mono

SSK

MOOK, VARIATION IN SPIRIFER MUCRONATUS 199

Spirifer mucronatus mut. alpenense—Continued

: Adult Stage Neanic Stage

Number ‘Width Height Index Index Width Bene “Index Index

(reverse) (reverse)

20) Sars 41mm. 20mm. _ .487 2.05 41mm. 12mm. .292 3.41

BS ova sie eae Stet Seats 6 af eA ears Apa Ses fap

BES cca ws 32 15.5 .484 2.06 32 10.5 328 3.04

73 6 Se ee Aan 4h ASAE eae BAe ay ae ate eet

2 30 16.5 .550 1.81 30 BLISS .383 2.60

BOG oc 31 pL ms) . 564 sla of 4 31 14 .451 2.21

24 oe

22 2) eee 31 15.5 500 2.00 31 9 290 3.44

2 5 en See

2) |e 35 23 657 1.52 35 14 400 2.50

Sn a. 5, sce es aes Bucs

= Van a 30 15.5 .516 1.93 30 8.5 . 283 3.52

BUS eae <.« 27 14 53s} 1.92 27 125 BAT 3.60

BUA ie = 35 20 aL 1S 35 12 .542 2.91

SOD... eee aoe ane sy -

BUG as 3s 30 LS 083 a ar a 30 ain 366 22

3; | 7 (ae 32 IG dol 1.88 on iS 390 2.56

= | 2 eee 39 20.5 525 1.90 39 12-5 320 Se 4

B09 es... 35 eps) . 648 1.55 Sees Bh

21 ae 25 Lars, . 700 1.42 25 9 . 360 PA Uy |

21 el Ea 34 23 .676 1.45 34 12.5 . 367 2.72

51 7 26 15.5 .596 1-67 26 9 .546 2.88

Se Ss 0 = 33 Pape . 666 1.50 33 12.5 318 2.62

31 2 Se Ghee

21 36 16 444 2.25 36 4 (a Se) 319 3.13

SNES Sais « eek

Nee ces 26 15 576 1.73 26 ich 423 2.36

Bac aie. ot 18 432 205 37 9 243 4.11

= sae

= 38 18 473 read Ia 38 10.5 276 3.62

ie... aoe ee sbe ae

3a 26 16 615 1-62 26 12 461 2.16

32-3 37 i) 513 1.94 37 13 348 2.84

Sd 32 27 843 dies i 32 15 .468 213

2) 26 12 461 2.16 26 C5 .280 3.46

BAG. 5 os ere ae

2 ae ol 18.5 696 aed 31 14 451 BrAl

SER. a's» ano. 219 .690 1.44 2S 105 .381 2.61

2 ae sara en

=o) See 38 19.5 513 1.94 38 12.5 .328 3.04

RE sais « 30 20 .666 1.50 30 12.5 .416 2.40

“2 Se Heihes ee ARS See Bisse « eae S eet

233 ae 25.5 14.5 .568 1.15 25 5 8 .o13 3.18

334...°..

3 26 as, 672 1.48 26 It .423 2.36

200 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. alpenense—Continued

Adult Stage Neanic Stage

Number Width Height Index Index Width Height Index Index

(reverse) (reverse)

331; See 29mm. 19.5mm..672 1.48 29mm. llmm._ .379 2.63

5 Se 29 ZA 25 741 1.34 29 13.5 .462 2.14

MMOs ses 29 20 . 689 1.45 29 13 448 2.21

Be Re aT 16 -592 1.68 27 $.5 514 3.64

=> eee 355 23 . 657 4252 3D 11.5 325 3.04

eo eee 32 17 531 1.88 a2 10.5 .324 3.04

o44..... 29.5 24 .813 4.22 yin} Pa 15.5 225 1:90

= ee 21 24 .888 1.122 27 14 918 12

aa hs ee 29 18 .620 1.61 29 11 ae pas

52! Hee ar : ff crate 2 ae

=) | Sees 28 18 642 1.55 28 10 oT 2.80

2] Pee ee 35 16 484 2.06 30 11 ano 3.00

5-9

3S eas ry f £2) . 159 fhe! a | 14 .518 1.92

222 Se oe Soak went ee ae Seite = aie = ahaa

= ee 54 19 908 1.78 s+ 9.5 .279 ope.

“3 eee hats See Looe —

as (ee eee 54 eae 661 41.51 34+ 14 411 2.42

ae)

[on

jet

oe)

ves

le)

04)

v4)

Nie

=e =e Do 19.5 .990 1.69 DD 15 393 2.53

5 29 17 . 586 £50 29 9 310 3.22

OG es. ae

50) ee 39.5 18.5 465 2.13 39.5 11 278 3.09

14.

15.:

7 6)

(oh)

Cr OT

Ye oon

co |

[eat]

~~)

Ww Ww

feet

on)

jet.

per

va)

& @

iD. =f = ee

5 ea 25 a 600 1.66 25 T .280 3.97

BAN oak 5 50 12.5 416 2.40 30 8 . 266 3.75

25 ae OL 20 645 1.55 31 14 .451 2.21

MD =e Sse 30 15.5 516 1.93 30 11.5 .383 2.60

SO. Se 27 16 a92 1.68 27 12.5 . 462 2.16

5 eee 21 20.5 976 1.02 21 11 .925 1.90

= ee Sigs eee Hi Aas ee ae nica

Se 3D 21 . 600 1.66 30 14.5 .414 2.41

oe ee 27 17 .629 1.58 27 11 407 2.45

B25 5.2 29 16.5 .568 1.75 29 415 .396 2.52

SG...’ - D1 18.5 .596 1.67 ol 11 .oo4 2.81

BOF tien we 25 16 . 640 1.56 25 10.5 .420 2.38

BBe: = Bo 20.5 .o81 1.70 30 12.5 .ooT 2.80-

= 9) 36 19 527 1.89 36 15 .o61 2.76

ore 36 15.5 .oT4 2.66 36 10 200 3.60

= | eee 29 19 653 1.52 29 11.5 396 2.52

Number

oseee

-eeee

“eres

ee ee

MOOK, VARIATION IN SPIRIFER MUCRONATUS

= .

aaa Stage

Height Index Index

(reverse)

19mm. .542 1.94

1'¢25 .486 pl

20.5 661 Bee |

15 . 656 1.53

1% 447 2123

16.5 ATI 212

15-5 . 645 1.54

19 . 703 1.42

19 .644 #.bS

Ay | . 700 1.42

IG 25 .500 2.00

17.5 .486 2205

16 .615 4 762

18.5 . 525 1.89

at .538 1.85

14 .518 e792

5 .968 4°76

19.5 590 1.69

1675 . 804 1.24

di . 700 1.42

15 550 1.80

20 769 130

18.5 . Et 1.40

20 . 689 1.46

2025 . 759 oof

i .625 1.60

20.5 745 ise b

20:55 745 gai Y

a1 750 De So

20 . T40 2235

2625 is 1.29

25.0 . 839 1.19

23 . 884 PENS

14.5 Sot 1.86

19 .716 1.39

21 807 4.23

18.5 .616 1:62

18 . 654 2352

20. 714 1.40

18.5 .714 1.40

rg 3 .811 1.20

18 . 120 7235

23.5 S70 1.14

DAS . 750 $35

a1 651 jas

Width

35mm.

Wo w w&

OmMmwWre DA

nis

bt |

21 Ww Ww Ww

bo w bo

CAAA to o&

WNwwd Ww ww wy wv Ww

CO 7a)

bb wy bw ty bw

ADDAA

or)

-~| -!

on)

Spirifer mucronatus mut. alpenense—Continued

Neanic Stage

Height Index

12.

13.

15.

10.

14.

Or Ot Ol

Or Cr St Ov Ov OT

OU Or

Or Ol Ol

Cr Cr

(reverse)

mm. .357

.355

.900

.413

apf 5

. 300

.458

. 444

.338

.4338

.333

.291

.4038

.414

384

.B70

.409

375

pt?

. 466

.dol

.480

201

ia?)

b> W bh

~~ Oo OO

rsAaes

WOWwWNNNNwWwH NWN WW bo

SS Ww Ww

ear

wm Wwe Wd Ww vw

=r)

NNWNrRNWrRFrRPRNWNNWPNNNWRPRPRPNY NR RNY WY Wb Wb

9()2 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. alpenense—Continued

seal SI se Neanic Stage

Number ‘Width Height Index Index Width Height Index Index

(reverse) (reverse)

AST we 29mm. 19.5mm. .672 1.48 29mm. 11.5mm..396 2.52

ASS. e ie 27 1OES . 709 1.38 27 9.5 .ooL 2.84

ASG Face 29 22 . 158 ese 29 aby - 586 1.70

440..... 30 19 - 633 ibe aye 30 14 .466 2.14

2 ou eae 32 20 .625 1.60 32 - 14 437 2.28

442;5..... 29 18 .620 168 29 10.5 :362 2ehG

443..... 29 1325 .465 2.14 29 10 344 2.90

444..... 22 af af: .500 2.00 22 9 .409 2.44

445..... 26.5 19 CLG ay 26.5 11.5 .433 2250)

446-468 inclusive = Spirifer mucronatus mut. thedfordense.

AGO. coe 32 20 .625 1.60 32 12.5 .390 2.56

AOS s Ge: 32 Pat . 656 UR 34 oe 13.5 421 Zak

cy Gi lee 28 20 .714 1.40 28 13 . 464 Fob

2 Se 28 26 928 L207 28 16 «tee eae

ANS sis 'ss 35 23 657 dio 35 16 .457 2.18

AWC. Ce 32 19 .593 a Br) Bye 1s . 390 2.56

ADF. ws 32 20 625 1.60 32 13 . 406 2.46

ANDO aie ens 29 .5 20 677 1.47 20-5 12 407 2.45

ca Ge ns 25 i ry 680 1.47 25 12 .480 2.08

AG SAS 6S 29.5 24 .814 a 29.5 15S .D08 1.96

BO 28.5 22 oreahel: 1.29 2560) 14.5 .508 1.96

ASOS seco. Sif 7A | OCT 1.47 31 13 419 2.38

ASTER ees 30 PAD . 700 1.42 30 13 433 2uou

Ae eos 2h 38 d Rot 486 2, 05) 38 ds 342 2.92

AGO. oe 3-9 Ws) 23 730 1-36 od es 12.5 .396 2.52

484..... 29.5 Dai ses 763 tee k 29.5 14 474 vee ht

7 oo ee 32 23 718 1.39 32 1055 Sat 3.04

486..... ree

PSH cere vers 36 oa 583 Lat 36 a he 333 3.00

BO elas states syst wows sks Reet

2 iy) eae BE 20.5 661 gers & 31 . 14.5 475 pope

490. oss. 26 Ze, . 846 1.18 26 Lb55 442 2.26

2 31 215 693 1.44 = AS 20 435 2.29

21 ee 31 17.5 564 a 7 a ¢ 31 125 403 2.48

2 eee PA ape 880 g eg 25 14 560 1.78

494..... 30 Lt5 583 Led 30 13:5 450 PAR

495..... ee seer awe Bs Sete <ul

A9Ge <<... 28 7d a 767 B25 28 LE ee 2.54

A ete une 26 A | 807 i War 26 a5 O17 1.72

498..... 24 23) Ba 895 a Ra ie 24 15 625 1.60

ae Jt 22 814 122 QT a By 407 2.44

is eee a 29 23 . 793 1.26 29 12 413 2.41

NT cass, ce 30 Dit. 700 1.42 30 13 433 2.30

21 ( ee 35 Pe 657 io 35 13.5 385 2. 59

MOOK, VARIATION IN SPIRIFER MUCRONATUS 203

Spirifer mucronatus mut. alpenense—Continued

Adult Stage Neanic Stage

Number “Width Height araniee Index Width Height _ Index Index

(reverse) (reverse)

BOa. «se 25mm. 19mm. .760 dS 3 25mm. 12mm. .480 2.08

m4... 33 24.5 742 1.34 3 15.5 .499 2242

= re 33.5 PA .626 1.59 33.5 10 .298 3.30

SOS 6 sss 2 Bas . 833 120 27 14 018 1.92

ess ees 36 Pe e3..565 Pee 36 1225 347 2.88

eee 30 23 . 766 1.30 30 10.5 .350 2.85

; | ae Se. do -) 696 1.43 28 10.5 375 2.66

2) 28 19.5 . 696 1.438 28 13.5 .482 2.07

Pi. re ss. 28 24 . 856 1.16 28 13 .464 2.15

PE = ae 25 21 . 840 eS 25 14 . 560 1.78

57 36 26.5 . 736 ESS 36 16.5 .458 2.18

Bey ns 28 pAb . 750 iss 28 13.5 .482 2.07

be ss < « 394 Zoo . 734 1.36 3 9 281 panies

BGs vss 30 rls . 700 1.42 30 9 300 Stas s

a 40 19 475 2310 40 aa | 275 3.63

AG Ie. < <= 30 LES .583 SEA. 30 10 333 3.00

TiN yee oa ee oes er Ectae Joi oF

=), ae 30 22 PY 33 1.36 30 pW 3) 416 2.40

5] 26 26 1.000 1.00 26 13.5 .519 1.92

= ae 26 15.5 .596 E66 26 8.5 .326 3.05

Gea sha. a 1525 .500 2.00 BH) 8 as 3.87

52 Se 29 22 Bf (3c LSE 29 12.5 431 2S2

5 Bears sotde rs ae Sahel at ie re soe

ip a5) 21.5 .614 1.62 35 10 . 285 3.9

Si ee D5 22, . 880 a ad 13° 25 9.5 . 380 2.63

Be see « Zt Ze Brel 1.28 27 12.5 .462 2.16

4 31 19 GE: 1.63 ok 10.5 308 2295

al) 28 21 .750 1.33 28 13 464 2.15

eye Ss a « 30 19 . 633 1.57 30 10.5 350 2.85

52 35 a .628 £259 35 ee 314 3.18

lacs ss 27 18.5 EGET: 1.45 27 ‘9 333 3.00

Bot... 29 19 .655 Pie 29 8 275 3.62

eis.» = 30 20.5 . 683 1.46 30 10 333 3.00

MO. sss. 24 A nS . 895 1 We | 24 14 583 Ea >

ae 26 ya . 865 #255 26 13 500 2.00

= ae 24 1925 .812 1.23 24 10.5 437 2.28

B59: .... 25 21 .840 1.19 25 13 520 1.92

J. 26 20 . 769 1.30 26 14 .576 1.85

Bes 5%. 0 26 18 .692 1.44 26 10 .384 2.60

ADs... 24 19 191 1.26 24 13 .500 2.00

Ree. es 24 19 erg ae 1.26 24 7 .291 3.42

TAT... ot. 24 .640 1.56 aie 12 320 3.12

MRSS cs = 23 19 .826 EAE 23 9.5 413 2.42

3 ae 24 17.5 . 129 1:31 24 10 416 2.40

24 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. alpenense—Continued

Adult Stage Neanic Stage

Number Width Height Index Index Wadi) SEapat Index Index

(reverse) (reverse)

> | a 28mm. 21.5mm. .767 £3 28mm. 13mm. .464 2.15

a5 ee tee 24 19 eed 1.26 24 12 . 900 2.00

Se 14 . 200 2.00 28 11 Ba 2.54

27 2 27 225 . 833 1.20 Zi 10 570 2.70

a ee Zo 18 .620 1.61 29 12 .415 2.41

25 Te 24 22 .916 1.09 24 13 541 1.$

3 | pee 34 aS .632 1.58 34 13.5 .o9T 2 oe

7s] Ces 27 19 . 703 1.42 2T 12 .H44 2.25

= 5 Were 28 19 .678 1.47 28 15 . 464 2.15

=" 7 | Te oes 31 21.5 .687 1.44 3 11 354 2.81

=f | eee = ZA . 900 ith 25 ct 440 2 27

Ses fhe 25 12.5 500 2.00 25 I 440 2.27

ME Se 2T (eae 648 1.54 at 11 407 2.45

esos se: 24 23 958 1.04 24 15 625 1.60

it ZO 20.5 TO6 1.41 29 14.5 500 2.00

= 3 ee oe pd | 677 1.47 31 14 .451 2 2F

2 ee 27 23 $51 erage 21 3.5 .500 2.00

= Ty ees 30 24 800 1.25 30 15 500 2.00

5 (, ee 30 20 666 1.50 30 13 .433 2.30

569 33 27 S18 1.2 35 14 .428 2 ae

Sy | oe 27 23 $51 1g rt | 16 .592 1.68

3 ft Soa 25 24 960 1.04 25 10 . 400 2.50

2 i eae 31 27 S70 1.14 a1 14 451 Zoe

ered =. - 28 20 714 1.40 28 14 .500 2.00

=. eee 28 20 714 1.40 28 14.5 51T 1.98

aris) 29 24 S827 ta 29 14 .4838 2.07

27) ae 25 20 S00 i 25 13.5 540 1.85

= 7 7 hee 30 22 i335 £3 30 13 .483 2.30

oT); Sa 26 22 S46 Ae We 26 14 538 1.8

if. er 26 14 538 1.85 26 11 425 2.36

=> | ee 28 18 .642 1.55 28 12 .428 Dae

— 5] er 30 18 600 1.66 30 11.5 .388 2.60

582 28 22.5 803 1.24 28 at .410 2.45

23 fees 31 16.5 doz 1.87 31 8 . 254 3.87

Jo: Soe 25 21 S40 1.19 25 12 .480 2.08

wep 36 22 611 1.63 36 13 361 2.76

is ee 28 22 .785 1.27 28 14.5 517 1.8

Ra Pece « 25 24 . 960 ay 25 14.5 . 604 1-72

> as Mae 27 19.5 . 722 1.38 27 12 444 2 25

3°. Papas e 27 17 .629 1.58 27 9 .333 3.00

| ener 19.5 13 . 666 1.50 19.5 7.5 .384 2.53

2. 1 ee 30.5 20.5 .672 1.48 30.5 12.5 .409 2.44

= 32 17.5 46 1.82 32 9.5 -296 3.36

<3 eee 28 23 .821 1.21 28 15 554 1.86

= a ee 27 15.5 OT 1.74 27 10.5 388 2.57

Number

MOOK, VARIATION IN SPIRIFER MUCRONATUS 205

Spirifer mucronatus mut. alpenense—Continued

Adult Stage Neanic Stage

Cyaan ene ides) ade Width Height Index Index

(reverse) (reverse)

25mm. 25mm. 1.000 1.00 25mm. 15mm. .600 1.66

35 g eet) aS Ty 2.80 35 9.5 al 4.10

33 22 . 666 150 33 13 .393 PES:

29.5 20 677 1.47 29.5 gi sake 2.68

24 15.5 .645 1.56 24 10 .416 2.40

28 20 714 1.40 28 13 . 464 2.15

28 18-5 . 660 IDK 28 5b .392 2.54

29.5 20 677 1.47 29)..5 gy es .3889 2.96

28.5 19.5 . 684 1.46 28.5 12 421 Beat

30 13.5 .450 De22 30 9 .300 S.oe

Spirifer mucronatus mut. profundus

23 15 .652 1.53 ae gS a .500 2.00

24 Ta35 .562 dP i 6 24 9.5 395 232

22, 155 TO4 1.41 22 gh les Rae ae

2a 18 666 1.50 At 13 481 2.07

25 17 680 1.47 25 9 360 PA

30 16 boo dea ov! 30 g te .366 2.72

28 15 535 1.86 28 10 Bes 7 2.80

23 16 695 1.438 2a 9 .391 2.55

27 15 555 1.80 27 pW 407 2.45

28 16 571 ge 28 ui! .392 2.54

23 15 652 1.53 23 a | ATS 2.09

22 19 863 1.16 22 15 .681 1.46

a | 14 666 1.50 BA aft 05 DAT 1.82

ae, Ur 681 1.46 ves 9 409 2.44

34 18 529 1.88 34 10 294 3.40

2205 th 493 2.04 Dey 8 .3B55 2.81

Ze 19 863 ga a 22 12 .545 1.83

25 20 800 2a 25 14 560 1.78

27 16 518 1.68 27 10 toke 2.70

19 13 684 1.46 19 B55 .500 2.00

a, 13 590 1.69 22 8.5 386 2.58

23 ay (aS 760 1S. 23 11 478 2.09

23 19.5 S47 5 aa 34 23 11 478 2.09

1905 14 aa a 1.39 19.5 12.5 641 1.56

19 19 1.000 1.00 19 10 526 1.90

26 18 692 1.44 26 10 384 2.60

20 a ET WN 875 1.14 20 9 450 2.22

ak 15.5 738 5 21 10.5 500 2.00

23 16 695 1.48 23 12 521 We a

20 15.5 574 1.74 at 10 370 2.70

206 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. profundus—Continued

Adult Stage Neanic Stage

Number Width Height Index Index Width Height Index Index

(reverse) (reverse)

834.... 20mm. 13mm. .650 1.53 20mm. S8mm._ .400 2.50

S52. as 15 m5 43) 1.86 28 10 37 2.80

S36;,.0.1728 13 . 650 1.53 20 9 .450 2.2m

S37. 24 16 666 1.50 24 9 .375 2.06

S38 29 16 dol 1.81 29 10 544 2.90

S39 19.5 13.5 692 1.42 19-5 8.5 .435 2.29

S40 20 i+ \ 1.42 20 ae 475 2.10

S41 18 12.5 694 1.44 18 S . 466 2.25

S42 25 14 560 1.78 De 9 .360 PAS iy é

$43.. 19 17 . 894 1.11 19 11 018 ice

S44... 23 15.5 .673 1.48 = 9 47 2.55

4 San See 15 .625 1.60 24 9 .3T5 2.66

S46.. 27 13.5 . 900 2.00 27 9.5 sol 2.80

7 : eee Sake 2h

Ns | Se 15 S57 1.16 47 5 9 514 1.94

S49 Za 14 482 2.07 29 10 344 2.90

S50 30 15.5 516 1.93 30 9.5 316 3.15

rt ee 16 592 1.68 27 10.5 388 2.51

852. cera aere 2 aa ze

5 5 fee a et 12.5 694 1.44 18 rf .388 2.57

S54 19 10 526 1.90 19 6.5 .342 2.90

Sy 15 600 1.66 25 9 .360 2.7T

S56 24 15 625 1.60 24 9 .3T5 2.66

S57 27 14 518 1.92 27 9.5 dol 2.08

BS =. 225.5 18 . 705 1.41 25.5 10.5 411 2.42

S59 19.5 13.5 692 1.44 19.5 8 .410 2.68

S6U 24 15 625 1.60 24 10 416 2.40

S61 23 16 695 1.43 23 10.5 .456 2.19

SR a a 11 500 2.00 22 6.5 2 3.38

2 = 15.5 673 1.48 23 9 391 2.55

aot... B25 13.5 600 1.66 7 ae 9.5 .422 2.36

Mir. . ... SD 17 S50 gs as 20 9 .450 y Sip a

S66 23 13 565 AK ic | 23 9 Be 1D rT

S67 22 16.5 750 1.33 22 11 500 2.00

S68... 20 12 600 1.66 20 8 400 2.50

3 Ee ohh : wants

|| ee Pe 11 468 aes a 8 340 2.93

ye ghe nose BF ee Pie Big oa

72 24 14 583 cy I 24 9 .3T5 2.66

Nachos Ve | oe 16.5 6ST 1.45 24 10 .416 2.40

S74 28 17 7 1.64 28 10 oot 2.80

Pics ee 15.5 7 1.41 22 9 .409 2.44

S76. a n a

S77 23.5 15.5 659 it 25.5 9 382 2.61

i te ee 23 12 o21 1.91 23 s 347 2.87

Number

STD. «

21,0 eee

SOPs sis.

313 ae

BO ca

S84...

Bons swe

po ae

So so.

Boa. .:.

893...

94...

BOR...

S96. ...

Ber...

2 ee

Bao).

900....

901....

902...

903...

904....

905...

S06... -

Sie...

908...

909....

Bt... 2).

iS.

Bi...

915....

SiG...

Bay...

ate. .

m9...

920...

oS...

925...

MOOK, VARIATION IN SPIRIFER MUCRONATUS 207

Spirifer mucronatus mut. profundus—Continued

Adult Stage Neanic Stage

Width Height Index Index Width Height Index Index

(reverse) (reverse)

20mm. 14mm. .700 42 20mm. 8mm. .400 2.50

24 18 . 790 feos 24 8.5 O04 2.82

26 20:5 . 788 1.26 26 10.5 -403 2.47

yA § 16.5 785 227 21 10 -476 2 AD

1825 14.5 chef I beer £825 10.5 567 176

ah 15 .714 1.40 oe 9 .428 eves

28 AG 607 1.64 28 11 892 2.54

22, 1 045 hess De, 8.5 086 2.58

24 tae . 645 £62 24 9 wee 2.66

BA 16.5 660 £51k 25 9 .360 APE

2A 16 666 1.50 24 9 a 2.66

20 tans 675 1.48 20 8.5 425 yes

ei! 14 666 1-50 A 9 .428 Zhe

20 a | 650 £81 20 F sou 2.85

29.5 16 627 1.59 25.5 10 392 2.55

pe NDS g bye 489 2.04 24.5 9 . 367 pe ye

24 16 666 2506 24 10.5 ee Syl 2.24

ia Paes 15 697 1.43 wey io 348 2.86

18 16 858 ee 18 9.5 527 1.89

16 15 843 £uls 16 7 437 pad

16 13.5 843 LA8 16 8 500 2.00

28 16 yr eS 28 10.5 375 2.66

EI 14.5 676 1.47 LF 8- 470 2. 32

2a 17 755 #232 wee 10 444 2.25

20 13:5 675 1.48 20 8 400 2.50

17 EE 647 155 17 8 470 2.12

18 A ay 694 1.44 18 8 .466 pe 5

19.5 14 fay £39 19.5 7.5 384 2.60

2 14 608 1.64 = 8.5 3869 22tO

19 15 710 1.40 19 8.5 447 2.02

20.5 AES 814 dia 7) 20.5 8 390 2.56

19 13 631 1256 19 as 384 2.60

P.5 14 800 $25 VF-5 8 457 2.18

as Be Bat 1.91 23 9 347 2.55

18 fs 750 Se 18 8 444 2 25

eee

ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. profundus—Continued

Adult Stage

———

Width Height Index Index

(reverse)

2imm. 13mm. _ .619 1eGL

25 13 .520 1.92

19 15 .673 1.26

27 20 . 740 1.35

AT. 5 15 . 857 1.16

17 14 . 823 fe21

19 15.5 .815 4.22

23 12.5 .548 1.84

21 15 .714 1.40

30 20 . 666 1.50

24 10.5 .479 2.28

35 14.5 414 2.51

18 3 hee 1.38

20 14.5 125 1S

18 11.5 638 1.56

24 14 583 ier gb

24 14.5 604 1.64

22, 13 590 1.69

18 12 666 1.50

15 12 S00 1.25

pal 14 666 1.50

27 19.5 722 1-37

21.5 14 651 1.53

19 12 631 45S

20 12 600 1.66

16 10 625 1.60

22 17 172 1.29

18 12 666 1.50

19 14.5 763 a hes 3

18.5 15 818 2s

21 15 714 1.40

22.5 16.5 Tre 1.36

eal | 16 T61 1.31

19 15.5 815 £22

18 15 . 838 1.20

25 16 . 640 1.56

22 12 545 1.83

21 14 666 1.50

2A .5 13 604 1.65

18 3 722 1.3

18.5 10.5 567 1.76

21 14 666 1.50

a5 12 .631 1.58

Width

21mm.

21.

Nmomwh:

IND -

Neanic Stage

Smm.

=~] ©

“1 ©

fj.

— O—

Conmwwom ds wo

OO 00 OO

“1 00 ©

-1 =~] =I 00

=1 00 0

Height

Cr Ol

i |

Index

(reverse)

.380

.320

.421

.310

.514

411

AAT

.369

.do2

.300

.309

.228

.416

.425

.388

291

.375

.318

.961

.500

.380

.425

372

.421

es ¥0s)

. 468

ATT

.444

Index

Doe:

WDHHENNNNYNNNNNYNYNNNWANWNHNNHKRWNNYNNNRFNYNW

wNWwWrRNMNN:

.62

.12

Number

DIA.

OPO... 0.

oy eae

DTS... s

2 ee

US...

Des S:.

982...

DOS ao e's

984....

985

988... .

Jo re

2 ae

OOD...

998...

929... ...

TOGO... .

MOOK, VARIATION IN SPIRIFER MUCRONATUS

Spirifer mucronatus mut. profundus—Continued

Adult Stage

(Cae e « hin aay

Width Height Index Index

(reverse)

25mm. 16mm. .640 1.56

Ze ay 15 . 697 1.43

22 a5 022 LOT

18 AS Pl (e 1.38

15 9 . 600 1.66

18 ps (22 1.38

18.5 12 . 643 1.54

20 2.5 . 625 1.60

17.5 12 .685 1.45

29 a2 .413 2.41

18 dl . 638 1.56

21.5 P25 . 720 1.38

17 bce 647 1.54

19 14 736 £35

25 16 . 640 1.56

19 13 . 684 1.46

22 16 120 1.37

25 16 744 1.34

23... 5 18 765 230

29 14 .482 Lert

20 m5 . 625 1.60

22.5 14 622 1.60

35 18 .o14 1.94

28 18.5 . 660 “2.51

23 18 . 782 1.28

20 19 . 950 1.05

pepe 13.5 .613 ih: G2

25.5 18 .113 1.41

20 14 . 700 1.42

25.5 16 627 1.59

19 12 .631 1.38

19 Ly. . 894 ays 2 |

20 14.5 725 1.37

at 20 740 1.35

22 19 S63 HAS

22 18 .818 e222

20 i ite 600 1.66

25mm.

AA Wes

Ze,

Neanic Stage

eee

Width Height

10mm.

eee ee

Ol >

“141

OL OT OL OI

co |

Index

(reverse)

. 400

. 465

. 980

.444

.033

.416

-432

475

428

.258

.416

.488

441

447

No +

bo «

Ww 1

NWWWw: HB bwbd-

bob bo w

NW WwW -

pe eee Pe Pe be bebe et Be Sto be ee bo be

210 ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. profundus—Continued

Adult Stage

—<—<—<———————

Number Width Height

1023.... 22mm. “18mm.

1024. 23. 26 18

AQ eae pA

1026.2 24 15

1020 eee 16

1028.... 22 16.5

1029. ou 2 18

10 Faas 15

4035 Jee Se 16

1036.... 238 13

IOS io. Gee 12

1038.. 28 16.5

1039... 18 10

1040.. 20 15

1041 19.5 42.5

1042 24.5 19

1043. 20 13.5

1044. 21 16

1S 2S 16

1046.. 25 15

1047.. 28 14

1048... = 16.5

1049.. 21 3

1050 18 14.5

1051 19 13

1952. 35. 48 12

1053 18 13

1054 20.5 125

1055 eS

1056 22 18

1057 23 16.5

1058... os ae

1059. 35 14

1060.. | 16

1061.. 25 16

1062 20.5 14

TGS 55 Be 18

1064 24 16

1065. at 15.5

1066 29 15.5

1067 26 13.5

1068.... 25 21

1GS 2 235) Soe es

1070.... 28 14.5

Jot

v4)

IOTL 2s Peace

_ Index

(reverse)

.818

. 692

.954

.625

961

. 750

. 720

. 833

727

565

666

589

B55

750

641

175

675

761

5T1

A fe ek RA BA RE

ll coal ll oll ll ll al ol ol ll ol ool ol oll el wel lo

Stl oe sell ll coll ell oll wll ol ool oo

Index

22mm,

“Width

OV Ol

Wh.

WNNONNWNWhHyNMNWwh hd bd .

HAHHAASA HAS & i

=~] Ol

Neanic Stage

Height Index Index

(reverse)

12mm. .545 1.83

11 .426 2.36

10 .454 2.20

9 .375 2.66

9.5 .333 3.00

10 » .454 2.20

mks .440 2.27

10.5 .583 ae.

10 .454 2.20

8 347 2.87

7 .389 2.59

10.5 .875 2.66

7.5 .416 2.40

8.5 .425 1.35

9.5 487 2.05

12 .489 2.04

10 .500 2.00

10.5 .500 2.00

pitt - 392 2.54

12 .480 2.09

5 .3821 3:44:

10.5 .456 2.19

.428 2.66

9 .500 2.00

8 .421 2.37

8.5 .461 244

10 .555 1.80

9 .489 2.27

10.5 ATT 2.07

12 521 ie.

11 .o14 3.18

10.5 .388 2.57

10.5 .420 2.38

9 .439 220

10.5 .338 2.95

11 .458 paw §

i 291 3.42

9 .310 S22

9 .346 2.88

id .440 2.2

8 AT 2.87

11 .392 2.54

11 .415 2.40

Number

BOT 5:2.

1073

HOT4:....

OT si

ONG. .:

Oe Grane

BOTS 6 3 x:

4079...

FOSO.... .

1 od Gee

1082

AOI:

BARON

HOA.

Ts...

POG. 3.

qd07...

es:

1109...

ttt)...

Jo es

ie Oa

ide aoe

mG...

ay: .

OD...

wia0 3. .

i a

HAS. a.

1124....

WE...

E26...

HIS... .

iiss ee

MAG 20s

MeN ese <\.< «

HiS4...

m135)....

MOOK, VARIATION IN SPIRIFER MUCRONATUS

Spirifer mucronatus mut. profundus—Continued

edule Stage

Width@tGeient . Indes - Tndes

(reverse)

29mm. 138.5mm. .465 2.14

20 13 .650 tbs

Da 2, Dao MEST

26 Ae .461 216

28 20 114 1.40

26 * 14.5 a5 506 79

24 153) . 562 deri

28 15 535 1.86

26 Ly .653 fe 52

24 18 . 750 hos

20 iS. oF 675 1.48

21 125 595 1.68

22, $25 . 795 1.28

oT 16.5 al 1.63

21 15 .714 1.40

a af 681 1.46

32 20 “625 1.60

26.5 18 .679 1.47

41 14 651 eae:

PBS $5.5 674 1467

24 ale¢ . 708 1.41

a2, 17 (te 1.29

MAYA, 16.5 . 702 1.42

2.0 19 . 744 1.34

20 16 . 800 a Leet

26 14.5 ave eo.

Pare LE .629 1.58

21 15 114 1.40

Pai 19 . 904 1200

22 a es . 195 ame

1 ef mss . 882 ole

aah 15 . 714 1.40

ak ie .619 6k

20.5 15 fou 1.36

34 13 541 1.84

ay Pie L029 OF

28 17 . 607 1.64

24.5 13 .530 1.88

Width

29mm,

on:

Neanic Stage

Height Index

(reverse)

7mm. .241

9 . 450

8.5 £51 if

9 .346

a la a) .410

9.5 .365

10 .416

955 .365

at .423

gla 8 .458

9 .450

9 .428

11 .500

alas 407

10 .476

10 .454

12 375

11 .415

10 -465

11 478

dia .468

10 .454

dit .489

A 470 .

9 .450

9 .346

12 444

9 .428

10 .476

9.5 .431

9 .529

11 .523

8 .3880

8 .381

9 375

9.5 .556

13.5 .446

8 326

CO |:

ws.

Io bd bb

NONONNWNNNWWDhW-

a .

(op)

BFNNNNNNNDW-

wd)

Ne bw bo

=~]

19 4)

caw

jen

—— "“*s**

ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. profundus—Continued

Adult Siage

Width

18.5mm.14

8.5 15

22 1b

18 gh

~~)

ier

hol

ie)

Height

Or Ol

Index

(reverse)

.omm..

789

. $16

OV OT

fob fot

lO = °

fuk fed fh oh sk fk fh eek

fend fom fo fest fk

I beh et LD

feat feok fo

Juhl otek fk

Width

G Go dy

IN ¢

Pan)

yw ‘

BEEBEBBA

Neanic Stage

Height

cronatus mut. multiplicatus

12.

. Smm.

Or Ot

or on

Or Or

Index

(reverse)

.452

013

wp FF

ve)

RHASwRHSw: py

. . . om lo to eg

PPA GRRSMERRROE: ERR BE: Ses

WhNwWNWhNW NNW wh hb w:

lobo:

WSREBA: BS

WWW wh:

Number

me nee

MOOK, VARIATION IN SPIRIFER MUCRONATUS

Spirifer mucronatus mut. attenuatus

Adult Stage

Width Height Index Index

(reverse)

26mm. 15mm. N76 Lee

2D 15.5 .620 1.61

42.5 15.5 . 364 2.74

31 3 .419 2.38

24 a IF Gar G25 1.37

30 15 .500 2.00

28 15 .535 1.86

28 14 .500 2200

24 14 583 tk

28 1a 6 ST 1.98

19 12 .631 1.58

AT 14 295 3.35

30 13 .433 2.30

26.5 12.5 4AT1 242,

3 15 .500 2.00

30 8 . 266 3.75

23.5 ple: .468 ZAS

19 14 3 1.35

20.5 13.5 .658 1.51

29 15 LT 1.93

35.5 15 422 2.36

26 12 .461 2.16

32 12.5 390 2.56

25.5 12, .470 72,8 Ly

29 14.5 .500 2.00

26 AZ .461 2.16

2S 9.5 .413 2.42

28 14 .500 2.00

26 11 407 2.36

25 12 .480 2.08

37 14.5 ook 2.59

34 16 .470 212

23 13 .965 1.76

20.5 12.5 . 609 1.64

30 11.5 .383 2.60

24 13.5 .562 sary

29 11.5 396 2.52

——

Width

26mm.

LO s

ore

ee)

Dd bo wy Ny w

OU bo

Neanic Stage

—

Height Index

(reverse)

8.5mm. .326

10 . 400

10.5 247

9 .290

10 . 416

10 .333

10.5 375

10 Oot

9 mY (5)

10.5 315

8 .421

10 Bea

10 .333

9 339

9.5 .316

6.5 .216

8 . 340

8.5 447

8.5 .414

11 soto

9.5 267

8.5 326

9.5 . 296

5 . 294

10 344

9 .230

7 . 304

8 . 283

i . 269

S25 340

iE) .297

12 .358

¢ . 504

8 .390

8.5 .283

10 .416

7.5 258

213

Qo »

we)

Yo»

[td bs) .

WNNWNNMs NWN WW

IO Go He

Ls Oe Woo)

IO G

Co OO

; Index

.05

<> Op he

CO bet CO Co

=I oO

—

oa w

oS GB Os

Do he

A Oo Ww

raw)

fomed

. Number

ANNALS NEW YORK ACADEMY OF SCIENCES

Spirifer mucronatus mut. attenuatus—Continued

Adult Stage

Width Height

33mm. 10mm.

36 15.5

24 14

35 17

33 15

31 17

44 16

36 14

36 13

31 13.5

32 11

31 175

25 12

32 3

30 13.5

32 14

35 13.5

34 14

42 15

34 17

28 12.5

Sb AG

35 15

29 13

26 12

25 13

34 14

23 13

35 15

24 11.5

26 12

39.5 15.5

29 12

27 12

25 9

28 11

29 13.5

Index

(reverse)

.303

. 430

.983

.485

454

.948

963

.388

361

.435

.343

. 564

.480

. 406

.450

437

.385

411

oot

.500

.446

.D07

441

eat

NMwWWwhNM:

Www eR bh

NUON bos

NrRMYNYNW Wh:

Nowe Ne

Ooh w bw Wb bv

Width

33mm.

ie)

Lo ee

OV =I

Neanic Stage

Height Index

(reverse)

7.5mm, .227

10 <ahe

9 31d

11 .314

9.5 200

12.5 .403

10.5 . 238

ie! . 805

9 . 250

9 . 290

Fie: . 234

AS | . 354

8.5 340

10 .812

9.5 O16

9 .281

10 . 285

9 . 264

9.5 . 226

11.5 .838

8.5 . 303

a Be Be 2 . 365

10 .291

9.5 oot

7 . 269

8.5 .340

9.5 .279

co .326

9 257

8 es:

8.5 -326

9 227

8 275

‘Ge 2h

6.5 . 260

7.5 . 267

8.5 . 293

Index

Nr WHR wR DY ww WY

wWowwokr-:

wWwonNnwnr ww:

SeSAR

wwwwow eb w www wh Ww Ow :

RSS eS

.73

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_ care of

American Museum of Natural History,

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PRESS OF JUDD & DETWEILER, INC., WASHINGTON, D. C,

ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXVI, pp. 215-315

Editor, Epmunp Otis Hovey

REVIEW OF THE

PLEISTOCENE OF EUROPE, ASIA AND

| ~NORTHERN AFRICA

Henry FAIRFIELD OSBORN

NEW YORK

PUBLISHED BY THE ACADEMY

30 Juzy, 1915

THE NEW YORK ACADEMY OF SCIENCES

(LycEum or Natural History, 1817-1876)

OFFICERS, 1915

President—Grorer FREDERICK Kunz, 601 West 110th Street

Vice-Presidents—CHARLES P. BERKEY, RAYMOND C. OSBURN, — ~

CHARLES BASKERVILLE, CLARK WISSLER _ . ae

Corresponding Secretary—HENry E. Crampton, American Museum

Recording Secretary—EpMunD Otis Hovey, American Museum

Treasurer—HENRY J. COCHRAN, 389 Fifth Avenue

LIibrarian—RaLpeu W. Tower, American Museum _ a Ses

Editor—EpMuND Otis Hovey, American Museum aor

SECTION OF GHOLOGY AND MINERALOGY

Chairman—Cuar.es P. BERKEY, Columbia University

Secretary—D. W. Jounson, Columbia University

SECTION OF BIOLOGY

Chairman—RAYMOND C. OSBURN, 557 West 124th Street

Secretary—WiIL1i4AM K. Gregory, American. Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

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_Chairman—CLark WISSLER, American Museum

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The sessions of the Academy are held on Monday evenings at 8:15

o’clock from October to May, inclusive, at the American Museum of

Natural History, 77th Street and Central Park, West.

[ANNALS N. Y. Acap. Scr., Vol. XXVI, pp. 215-315. 30 July, 1915]

REVIEW OF THE PLEISTOCENE OF EUROPE, ASIA AND

NORTHERN AFRICA?

By Henry FAIRFIELD OSBORN

(Presented by title before the Academy, 12 April, 1975)

CONTENTS

4 Page

MUNIN IEM MENT ere tera ak Os ccc tuck erat ar tom eres ashe deel e, aud awe Wile oa canuhe mkarahanene Mee thts 217

Peete iaunal phase—_Quaternary.. i... 2. cc aes eee te becca sect eee ee ces 218

Means of establishing the time divisions of the Quaternary...,....... 218

Pliocene and Pleistocene life of Asia and North Africa........... piteletein st 219

SLES S O12 ASIEN A eet ae eek ee oe ee a a re eee ne eee. See lal 219

ee Me NCO Tab BINGE LCA c tatny fo ok Wintel oo nse tnt \aky dancin“: aadvevale! aribicyosase 4 wheda ciate 223

agra Sean AGTOME Ns coors oats, Ee Wa trae CR Sacra grate so-Gh0 Gri Be wtele bu See ree 225

Sources of the Pleistocene African life...... Uva aland ava add ste CNR ars 225

PRS aete ANG BTLOPCAI: AHIMITIES =< S59. a axnietd nS cue « o> chara ere: oyp @retd ele drale 226

African—Huropean distribution.............. tei avaca atelitla nth tar Ha ieee 228

Africa in Paleolithic and Neolithic times....... Gada Sui acne. aos.» oa CLS EP 229

Pleistocene or Glacial epoch in Europe.............. ins bees died shake ee 229

Muranon of the Pleistocene. ... 02... 00. cee eee Cphelavave: «tales wuatetintoretaraiees 232

PEN MUTT MIA Os TATAIN. o8Scrciciw betes g wie Sa'e arava swiss @ Soravet el oPday eee eraneee ene 235

Peete eeu Or the CUITUNE SHALES «26. 6 6 ose es oltcce ere eee wlereie sé Male oc ene 236

Hlevation and subsidence of land in Quaternary times.............. Zot

MRT MME SHI DIST CL ETHER ass otic ie chal ster ciate a utube; laa sepals Sioa us. x hae Biles wees OMAR 239

eMC TICE ToL Isl CRTs. s (eect tayratcrs tafe ay cre ae ede Wiis -s) ahs oases iarecy. Wie weeks «ere eye cater Obs 240

Pranic OF ClMAte AMG OTA... o6enies sad es os Sees at euidepe os oe ue Sen 240

Miavmetacial SLATE TLEMPCTALUITES:. 66s: cna co ce wee weiss eicle Seles wsleties 240

Moderate estimates Of temperatures... 26h. swe ce Gere we fe Ske wl 241

Warm and temperate interglacial Stages. .... . 66. dv ees cece oe cece 241

Perea ARON S VOL MOPAs 6< cdc. a o!2 acre(s. c. esi e 6 mie ee Sad aera Sak Mee: 242

RC ARO AOI OS Ol -PILRFOPE™ 5... ac sis oo wie eh au Rbadd be ws ei wesw ein eke O-6 ae 242

Me Sr reas, oy isha Seat hae: Eiaie © oe eho e hee oh Rd ee ib tee! 244

Eien Asiatic MAamMMals, WAIN FAUNA. sos woes ce tees ewe ce be 244

Kurasiatic forest and meadow, temperate fauna................ 245

Ee pEN TN STV ARTERIES. COM a feg MITUA ¢ Sitahang act: « wehawctaleel «, «od 40a, aialleha bivcaysnetea 248

ree ty Ole LISS oa “SIDERIA . a2) cs'cc's «ws oie ofa ees CUO 248

Hpndcavor Darren, Sround- Tama and MOTAs. . 2. -siccje ces see ece.e@ele 249

Penson tnenry Ol GOTas ANd faunas. . 6 cs .ie eases s die u's cece « oa eae dee 250

Le RVC OTH 29 J ea ceo OMIT AS Sa es RS 251

Pere LO EG er NET ELE Ls cieicis wit aic'st 6 even ine wt wale we Keeton aie stores wale aedid 251

+ Revision for German edition of the author’s work, ‘“The Age of Mammals ;’’ German

translator and editor, Dr. W. O. Dietrich, Kgl. Geologisch-paleontologisches Institut und

Museum, Berlin.

; Manuscript received by the Editor 25 January, 1915. (215)

216 ANNALS NEW YORK ACADEMY OF SCIENCES

Page

Culliire,stamessso0 6 ees: eee Aa es Stee eis eee So os Petes

Repetition of loess depositions.............. ene 2 LR

First Glacial stage—Giinz,. Scanian, NebraskKan...... tutes em (nie st « «oe

First Interglacial stage—Norfolkian, Aftonian, Giinz-Mindel............. 255

Humid. forest conditions 2/5 J2..0 oe ea Se nlemes wre oe wine ee 6 5 255

Wlera of the first: Interelacial) period: ss. 22 5252 4..5. a6. ee eee 256

Mammals of the first Interglacial stawe.....2 5.2.2. ... coy... eee

Hx GinchOns./s2 5 <5 96 6 =o eaten ei Stas & Siete ee a ae re

SUT VE VIS 2 sees es ise ns 5S a SP Sa a Sti ea en 257

Arrivals. <6. 0 ec ds oicde 5 ald coeten yates Seta eet me Saat ade oe ee

Wirst. Interglacial of Wranee.. 20.6 -6 2st 2. eee ee eee 260

Southern elephant (Elephas meridionalis) ............-...<cc+e--- 261

Hippopotamni. = 2.5.3 .eete vee ean Rhee eee ee bess ae nape ena 263

Sabre’ LOouhs «soa see ee eee ba wis Gis o ocdcls Se 263

Moose (Aicés) 20. Se. Ce A ee Oa eee eee 264

Problematic evidence of man... ..00....6. 02. se ike oo ee ee 264

Holithic theory : 2. cic Ses 6 Bone wie bee tle Se ee 265

Pithecanthropus: erecius. ts ce eee to oe wnt ae ee ee 266

Life of the Mediterranean islands.............. toe dala bad eee 267

Second Glacial stag Saxonian, Mindell, Kansan... ....2..)...4... See 268

Second Interglacial stage—Mindel-Riss.............. Frigates sina. , See 269

Moisture: followed. by aridity... 2.2322.2433 = 0k Se 3 eee 269

Climate. ~ os. 248 2 ows galdie dione ful vcm meabeieee ashe hare one 27C

Mammals. 2 9 oa sss ee dde od Soe sien. d deol ees See Re oe ee att

Survivals : :.2s <i ceeds woes skeen Seen kee ee 271

BixtinebiOns. « 2 ooie.cc isco. ee oc ies clones Se oa ale chee ee 272

APrival oo /.5.e0 Se SE RR bie ee SS ee re 272

Homo hewdeticergensisn. oo. 3. ee eee tue. d Sond ow oe 274

Chellean culture and ancient Interglacial fauna..................--- 276

Fauna of the Pyrenees, Cantabrian Alps, Spain and Portugal........ 276

Second and third Glacial and Interglacial epochs.... os bots ne ae ee 277

African—Asiatie Taman. 2h rs eke wteweie ds Cee een dais aie-e oo eet 277

Old elephant (Flephas ones) ere [ONE nites cts woe eee 277

RHINOCETOSES: ¢ ws ts SSE PE EE ee Ee 278

BovineS: 344 Js dene £2.60 Sc Seed dee bames cee eee 279

Megaceéros civantens.. 3225621. cek eee Ree eee mS ton nies ens 280

Hinphme or: ped deers. e232 oo 2siasne eae des vase ee cele See 280

Reindeer: 25... dasa eek Sen ee eee ona L's 'w.0 Soe cid cle ea 280

Carnivoress. 2.6086 20 Skea ee ee ee 281

Third Glacial stage—Hlinoian, Polandian, Riss........ gales be 6 ke oe 283

Third :glacial fauna. :.....)..2i0522) £20 Roc eke ee 283

Third Interglacial stage—Riss-Wiirm, Sangamon.................++-+++- 283

Climatic changes during the third Interglacial stage......... we ob ba ae

WOT as yj. ok oc 8S edela Nis « ie, cee Se Ri ee ee Ee 285

Wauna.. 22: 2.5 0t desican deeded es Jacana Oks de ee ee alee De ene 286

Piltdown man, Hoanthropus dawsoni............% cwule oe ees = on 286

Pre-Chellean and Chellean fauna. . 2... 2. sa. .2<smscec ccs eee 287

Chellean culture on the “low terraces”... ..... 0.0.6. 0s-0=-+-0 5 eee 288

Grays-Thurrock, Fiford (Essex, England).:....5.5..s< -. 22eene 288

OSBORN, REVIEW OF THE PLEISTOCENE 17

Page

Aeheulean Cultnre. fauna. sous oes Sew ea ee oe ree eee ee eee 289

WV ATTID SEAR C e oie oie ia ok wath ESL Gp eee ley ae ae 289

Cool Stase.4 se s5 oe PO oe eo oe ec ee ee 289

KRrapina Neagnderthaloid race! - oo Gen eo teas wo oe da eee eee 290

Mousterian culture, temperate fauna........ Ea AIOE SS Tay. drt ay! 290

I’ourth Glacial stage—Wtiirm, Mecklenburgian, Wisconsin.........,....... 291

Benning oc the remileer. and icave Periet.. io: och <2 0s oe ae motes ome 291

ering: OF the tmalVPlaCial WIPR I 25512 oso hiaace Se cca ae oie eee eee 291

Fauna of the fourth glacial stage........... SEM aed cele ee ae 292

BppeEr. Sands ior Mager ee kek chro ea ad ees sas wh Sie eae mae 294

MGS LEhian, T aAlLeOnenic: CUILUTE ; see oa le Se eee a ee eee 294

MSANG Eri Hal: PACES: Bd okie eneie «ure os <0 Se eke ori Ba ie seyes eched ener etapa rare 294

Postglacial stage—continuation of upper Paleolithic, reindeer or cave

ME ocr) ater tone ahs cS a canton ate wuss & ROE eae Bint 2 ia les aleharerm ate eee 296

MURRAY Rae Sr a tas MeN copie: a PN SteVaya an aE ral hoe ao we eh Oe 296

Mauser Pakecohthic, four. or five: human: races... 2.60060 /.« cA ewe we ee 297

Aurignacian, first upper Paleolithic culture stage................ 298

BermiEnedtle SECURE CULTURE. SLARGT Lo o2 Baca ls 2 sha Vow ess eae aoe 298

Magdalenian, third culture stage and fauna..................... 298

Ee MES noses ty eh OA eo cen WL sat ee 299

eM ETACTSIIIN EVE! ooo sacc,o Goo cleteck @ ciale «eine om PE ee rie 301

Re IE OAS oie Oo ears ele ee Gee oe etwas wie edie bine ee cS ORE 302

Migrations of the large mammals of the fourth glacial and _ post-

Seni TE RENEE ies PTR saee ee Pc, vie Gr vag El wre a aise Beet Weve CAS, wee os ae ae 303

2 t LSESG Hic PAAR IS SR mene re Pe or ie ge ae fa og Lee re A 304

Biber mptTINGe Cris ae Gar. Se FOE A So he Geol Sin Bias aia 305

i aOR EEC ae 20 Vere So ie «iar Goel os wetoiehelord ale k kie Sexton wR bust aw, aie oe ate ele 307

Horses of the Pleistocene................... i te oe eee ails 308

ere nes ee ONS Sige SON SD aoe nea Sat gre 310

Transition to the European forest stage......... eid, slenety, fie ey wnaisarte od, alalie 311

rete Heer ENE: EERO E A PA ©. ok bi. ciate alk GE ane mk Se ewe ede 311

ame ee eae BE I STEERS ESTAS (PA AITA EY, oso io Sau ve ors mls Se A BS De Seal ene we des 311

Survival of forest and meadow fauna........... Gace aerate eae aL2

Azilian-Tardenoisian, final upper Paleolithic culture............ ois

TIDES BTC 21 oe eae CC anne (a, SNe El ee So SRDS ee ceca ue 313

INTRODUCTION

We observe that the Upper Tertiary closes with a Pliocene northern

world rich with life, a world replete with Asiatic and African influence.

The Tertiary Period is followed by the Quaternary, or Age of Man, a

time of transition and of vast extinctions in Europe and North America

through natural causes, as well as of the geographic redistribution of life

and establishment of the modern zodgeographic regions. Toward the

close of the Quaternary Period man becomes the “destroying angel” and

very nearly completes the havoe which Nature has begun.

raw)

font

ANNALS NEW YORK ACADEMY OF SCIENCES

QUaLETHATY. oe onc emcee see Age of Man

WETHary Ft Skee ecm ee eee Age of Mammals

We thus enter a new Cenozoic faunal phase, the Seventh. When its

transitions are complete the world wears an entirely new and somewhat

impoverished aspect: the North has banished all the chief southerly

forms and established the five modern zodlogical regions of the Old and

New Worlds, namely: Palearctic, Nearctic. Oriental, Ethiopian, Neo-

tropical.

SEVENTH FAUNAL PHASE—QUATERNARY

IN THE NORTHERN HEMISPHERE THE GLACIAL EPOCH. VERY GRADUAL

EXTINCTION OR EXPULSION OF SOUTHERN TYPES OF AFRICAN, SOUTH

ASTATIC AND SOUTH AMERICAN ORIGIN FROM THE NORTHERN, OR

HOLARCTIC REALM. FIRST APPEARANCE IN CENTRAL EUROPE AND

NORTH AMERICA OF THE CIRCUMPOLAR TUNDRA FAUNA, IN EUROPE

OF THE STEPPE FAUNA. IN NORTH AMERICA EXTINCTION OF THE

REMAINING LARGE ENDEMIC QUADRUPEDS. THIRD AND FINAL MOD-

ERNIZATION OF EUROPE AND NORTH AMERICA BY THE HARDY FOREST,

MEADOW AND MOUNTAIN RUMINANTS AND THE CARNIVORES.

The grand geologic divisions of the Quaternary in the New and Old

Worlds are the same, namely, beginning with the Pleistocene and closing

with the Holocene.

( II. HoLocene. or RECENT EpocH. Mammals of prehistoric and recent

times. Domestication.

I. PLEISTOCENE, or GLACIAL EpocuH.

3. PoSTGLACIAL. Mammals of tundra and steppe type gradually

disappearing or retreating. Mammals of existing north tem-

perate type multiplying.

2. GLACIAL. Period of successive glacial advances and interglacial

retreats. Mammals of extinct and existing species com-

mingled. :

1. PREGLACIAL. Period of the lowering of temperature in the

northern hemisphere and modification of plant and animal

life. ;

(QUATERNARY

a ae

MEANS OF ESTABLISHING THE TIME DIVISIONS OF THE QUATERNARY

The fluctuations of climate and of the plant and animal life of the

Pleistocene are so numerous, so widespread, and so profound that it

seems best to introduce the subject by a review of the great time divisions

OSBORN, REVIEW OF THE PLEISTOCENE O15

Tod

ew)

together with some discussion as to the period when we should consider

that the Quaternary proper begins. The fullness and precision of Eu-

ropean faunistic investigation is in very strong contrast to the prelim-

inary results of American work, and in no other period may we anticipate

more weighty inductions from correlation between the history of the Old

and New Worlds. It is absolutely clear that a final and positive time

scale and subdivision of the early Age of Man are not far distant and

that the vast labors of European and American geologists, botanists,

zodlogists, paleontologists and anthropologists will finally be rewarded

with a harmonious theory of all the phenomena’ of the Quaternary

Period, the determination of the chronology of the various races and an

approximate estimate of the duration of the entire Quaternary Period

itself. The reader will observe that this correlation, derived from at

least five distinct branches of natural science, is based on evidence of four

kinds,

1. Geological: glacial deposits and erosions, which furnish the chief data for

estimates of time.

2. Botanical: plant deposits, alternations of northern, arctic, steppe, temperate

and southern floras, which furnish the chief data for estimates of tem-

perature.

3. Palzozodlogical: evolution and extinction of mammal and bird life, which

furnish the chief divisions of the Quaternary time scale and afford sup-

plementary knowledge of conditions of moisture, temperature and foresta-

tion. :

4. Anthropological: the successive stages of human culture and implements,

the skeletal remains of man, which combined furnish the minor sub-

divisions and correlations of Quaternary time.

PLIOCENE AND PLEISTOCENE LIFE oF ASIA AND NortTH AFRICA

LIFE OF ASIA

The region of the rich Tertiary flood plains of India? was one of the

main sources of the large mammals which wandered into northern Africa

and southern Europe in Pleistocene times; in other words, the large

mammals—the elephants, the rhinoceroses, the hippopotami—were all in-

vading forms from Asia and Africa. The relations between these three

geographic regions are, in fact, so close that they might be embraced in

a single zodgeographic realm were it not that throughout the Pleistocene

the forests and meadows of southern Europe also maintained a northern

Eurasiatic fauna which is entirely absent from southern Asia.

2See p. 323, English edition of ‘“‘The Age of Mammals.”

ANNALS NEW YORK ACADEMY OF SCIENCES

220

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OSBORN, REVIEW OF THE PLEISTOCENE

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929 ANNALS NEW YORK ACADEMY OF SCIENCES

The precise researches of Pilgrim,* published subsequent to the corre-

lation proposed by the author in 1910 in the Pliocene chapter of the

“Age of Mammals,’’* have resulted in a new classification and correlation

of the Tertiary flood-plain deposits of India which are of the utmost im-

portance and interest to students both of phylogeny and of geographic

distribution and migration.

In geologic time the Indian series extends from the Bugti beds, which

are of Aquitanian or Upper Oligocene age, to the uppermost Pliocene of

the Upper Siwaliks. The correlation with the successive intermediate

phases of European life appears to be quite close.

As regards the origin of the Proboscidea, the author discovers in the

Upper Oligocene of India animals which he believes resemble the Lower

Oligocene Paleomastodon as well as the Meritherium of the Faytim de-

posits of northern Egypt. These animals are referred to respectively as

Hemimastodon and Meritherwum. This discovery would favor the hy-

pothesis that the Proboscidea may have originated in southern Asia

rather than in Africa. Pilgrim believes that the Mastodon cautleyi of

the Upper Miocene of Perim Island gave rise to the Stegodon types of

the uppermost Miocene of Pikermi age, namely, to S. cliftc and S. bom-

bifrons, from which originated the genus Hlephas which first appears in

the dominant type Hlephas planifrons of Middle to Upper Pliocene times

in strata 3,000 feet above those in which the earliest forms of Stegodon

occur. :

The species H. planifrons is especially important because it has re-

cently been recognized by Schlesinger® in the Pliocene “Belvederschot- .

ter” north of Dobermannsdorf near Vienna. The horizon is regarded as

of Middle Pliocene or even earlier age. Pavlow® has also recorded the

occurrence of FH. planifrons from beds in Bessarabia which are regarded

as of Lower Pliocene age. In the Upper Pliocene of Europe occurs

Elephas meridionalis which is regarded as a descendant of L. pianifrons,

while in the Upper Phlocene of India occurs the Hlephas hysudricus,

which Pohlig considers as a geographic variety of the European H. merid-

ionalis. In the uppermost Pliocene of India also occurs the Dicerorhinus

platyrhinus, which is believed to be closely related to the D. etruscus of

the Upper Pliocene of the Val (Arno of Italy.

Some authors mistakenly regard the “Altelephant” (FH. antiquus) of

8’ PILGRIM, Guy E.: “The Correlation of the Siwaliks with Mammal Horizons of Eu-

rope.” Records, Geol. Surv. India, Vol. xliii, Part 4, pp. 264-326, Pl. 26. 1913.

+See Life of Southern Asia, pp. 323-332, “Age of Mammals.”

5 SCHLESINGZR, PAUL: “Studien tiber die Stammesgeschichte der Proboscidier,’”’ Jahrb.

d. k. Geol. Reichs., Vol. 62, pp. 87-182. Vienna, 1912.

6 PavLow, MARIE: “Les éléphants posttertiaires de diverses localités en Russie,’’ Ann.

géol, et mineralog. de la Russie, vol. xi, pp. 171-174. Moscou, 1910.

OSBORN, REVIEW OF THE PLEISTOCENE 2993

the European Pleistocene as related to H. hysudricus of India, but Pil-

grim and Pohlig rightly compare FL. antiquus with the Narbada elephant

(#. namadicus), which first occurs in the Pleistocene of Asia. In fact,

it is not known whether the phylum of F#. antiquus, which is quite dis-

tinct from that of FE. plamfrons-H. Meridionalis, originated in Asia or

in Africa.

To sum up, among the contributions of southern Asia to the Pliocene

and Pleistocene fauna of Europe are the following:

Elephas planifrons, entering Europe in the Pliocene, related to the Hlephas

meridionalis, the southern elephant.

EKlephas hysudricus.

Hippopotamus, H. javadicus, related to H. major.

Bison sivalensis, the short-skulled bison, related to Bison priscus.

The long-skulled Leptobos, related to the ZL. etruscus of the Val d’Arno and

Bos primigenius of the First Interglacial Stage.

The Sumatran type of rhinoceros, Dicerorhinus platyrhinus, related to D.

etruscus and D. merckii.

The hyzenas, related to H. crocuta, the spotted hyzena, and H. striata, the

striped hyzena.

The horse, Equus sivalensis, related to the Arab, or desert type of Europe.

Among the mammals which did not find their way from Asia into

western Europe are the camels and the various giraffoids. The absence

of the antelopine members of the Bovide is also a very characteristic

feature of the Pleistocene of Europe as contrasted with their abundance

in Asia and their presence in diminished form and numbers in the Upper

Pliocene of Europe.

LIFE OF NORTH AFRICA

It would appear that in Lower Pleistocene times when there were

broad land connections between Europe and Africa the latter continent

contributed to Europe some of its indigenous mammals and others which

had been derived originally from Asia. It is natural to suppose that the

hyena and hippopotamus, now so characteristic of Africa, entered Eu-

rope either from Asia or from the north African region. With these ani-

mals may have come the lion (Felis leo) and the “old elephant” (£.

antiquus), which is a primitive offshoot of the same stock that gave rise

to the African elephant (Lozxodon africanus).

We observe that in Lower Pleistocene times north Africa is still dis-

tinctively a part of the Ethiopian Region, closely connected with central

and southern Africa in its fauna. Throughout the Lower Quaternary

the fauna of north Africa is also closely related to that of Asia. More-

over it has a number of species in common with the Quaternary fauna of

DOA ANNALS NEW YORK ACADEMY OF SCIENCES

Europe. including those noted below which came into Europe from

Africa. The contrary theory of the relative geographic isolation of

Africa and Europe in Quaternary times originated with Pomel’ as the

result of his exhaustive review of the entire fauna of north Africa. He

concludes that since the resemblances between the European and north

African faunas are rare and often doubtful, the two continents were

for long periods separated by the Mediterranean Sea and Siraits of

Gibraltar.

VER UT

PLEISTOCENE

Fig. 1.—Pleistocene, or Ice Age

A period of maximum toial elevation facilitating free migrations and invasions of life.

culminating in the Glacial epoch, and followed by a prolonged depression. Portions of

northern Europe and the coasts of North America greatly depressed. Then a period of

reelevation. Rearranged aiter W. D. Matthew. 1960S.

Climate-—At the beginning of the Quaternary Period north Africa

was characterized by abundant rainfall which led to the formation of

great alluvial or flood-plain depositions. Im the Barbary and Sahara

egions the life was closely similar to the grand plateau life of equatorial

[oe

jay)

Lar |

faa)

—

(a8)

-t

ben |

fae)

Ler |

“

ay!

wild asses, giraffes, wild cattle, buffalo, antelopes. gazelles, gnus, elands,

hippopotami. wart-hogs, lions and hyenas. The presence of these ani-

7 PomeEL, A.: “Les Eléphants Quaternaires.” Carte Géol. Algérie, Paléont. Monogr.

Algiers. 1895.

OSBORN, REVIEW OF THE PLEISTOCENE 925

mals is consistent with the climatic theory of subtropical temperature

and alternate dry and rainy seasons.

Various phenomena point to increasingly long periods of drought and

progressive secular desiccation of this great region as the Pleistocene

advanced, resulting in the partial extinction and partial migration of the

great equatorial life into central and southern Africa.

Eurasiatic Invaston.—At the close of the Quaternary the bear (Ursus),

as a characteristic forest-dweller, requiring a moist climate, became ex-

tinct, while the Eurasiatic deer, wild sheep, wild boar, smaller mammals

of European type, survived and established for this region its present

affinity with Europe and its Palearctic fauna. We must account for

this northerly, or Eurasiatic, fauna of north Africa as having entered the

continent during the latter part of the Pleistocene Epoch and as sur-

viving in the forested regions of present and prehistoric times so as to

unite northern Africa closely with modern and prehistoric Europe.

North Africa thus becomes a part of the Palearctic Region.

Thus in no region of the world have more profound changes occurred

during and since Pleistocene times than in Africa north of the Sahara

Desert. : |

Sources of the Pleistocene African Infe.—Ilt is premature to attempt

to ascertain the original sources of all the various members of this im-

posing assemblage of mammals. ‘There remains always a great element

of doubt which can be eliminated only by the discovery of the complete

Cenozoic history of Asia and Africa. It would appear probable from

our previous studies that the several continents contributed to the remote

original ancestry of the African fauna somewhat as follows:

Africa or Asia, elephants and mastodons.

Northern Eurasia, deer and bear, wild sheep, wild boar.

Southern Eurasia, wild cattle and buffalo.

North America or northern Eurasia, rhinoceroses. various Hquid@, camels.

The most comprehensive comparison of the fauna of Africa and Europe

is that of Stromer,* in which the entire fauna of Europe and north

Africa, including the Reptiha and Mammalia, is compared from Lower

Eocene to Pleistocene times. This author observes? that during the

middle period of the Tertiary the mammal fauna of southwestern Eu-

rope, western Asia, India to China, partook of the tropical or subtropical

8 STROMER, ERNST: “Uber die Bedeutung der fossilen Wirbeltiere Afrikas fiir die

Tiergeographie.””’ Verhandl. d. Deutsch. Zo6l. Gesellschaft, pp. 204-218. 1906.

®°STROMER, ERNST: “Die einstige Verbreitung afrikanischer Siiugetiere.”” Naturwis-

senschaftliche Wochenschrift, N. F.. X Band; der ganzen Reihe. XXVI Band, No. 51,

pp. 814-816. Dec. 17, 1911.

226 ANNALS NEW YORK ACADEMY OF SCIENCES

character of the African high plateau fauna, rich in antelopes, giraffes,

zebra-like ancestors of the horse, elephants, rhinoceroses, hyenas and

apes. Late in Diluvial times in HKurope numerous representatives of

what we now consider a tropical African fauna, including hippopotami,

lions, hyeenas and apes, were widely distributed.

Asiatic and European Affinities——The total assemblage of the Pleisto-

cene life of north Africa may be summarized as follows from Pomel:

Man (Homo, Late Pleistocene)

Mastodon (Early Pleistocene only)

Elephants (several species related

both to Loxoedon and to Elephas)

Rhinoceroses (two species of the

African, or Diceros type)

Hipparions, zebras and asses

Camels

Giraffes (Libytherium, Giraffa)

Wild cattle (Bos), three species

Buffalo (Bubalus)

Dwarf antelopes, gazelles, gnus, Oryx,

nagor. elands

Hippopotami

Wild boar (Sus)

Wart hogs (Phacocherus)

Lions (two cavern species)

Hyzenas (spotted and striped)

Jackals (Canis aureus), India

Macaques (of northern origin)

Deer (of the Cervus type,

species )

Bear (of the Helarctos group)

Wild sheep and goats (Ovis paleo-

tragus, O. promaza)

one

A noteworthy distinction between

north Africa and Europe is the sur-

vival in north Africa of the masto-

dons throughout early Pleistocene

times; also of several species of hip-

parions side by side on the plains of

Numidia with the early north Afri-

can horses*® or zebras. Both the

mastodons and the hipparions are

absent in the Pleistocene of Europe.

We may now review the life of

north Africa itself in Pleistocene

times. Six species of elephant

occur, including the mastodons, the

southern mammoth (£. meridiona-

lis), and the “old elephant” (L. an-

tiquus). The most characteristic

and widespread elephant (ZL. at/an-

ticus) belongs to the African sub-

genus Loxodon while differing from

the recent African elephant (L.

africanus) in several points. The

latter species only occurs in the recent deposits of the latest prehistoric

period.

Similarly the two species of rhinoceros (D. mauritanicus, D. subiner-

mis) resemble the modern African types, but there is nothing to indicate

the presence either of the modern African “black” (D. bicornis) or

“white” (D. simus) species.

Among the Pleistocene horses, in addition to the surviving hipparions

and the species (H. numidicus) related to the Val D’Arno type of Eu-

rope, there is a third species (H#. mauritanicus) which exhibits tooth

characters of the recent zebra.

10 BouLzn, M.: “Observations sur quelques Equidés Fossiles.”

Ser. 3, vol. xxvii, pp. 531-542. 1899.

Thus there is every reason to believe that

Bull. Soe. Géol. France,

OSBORN, REVIEW OF THE: PLEISTOCENE 997

in Pleistocene times ancestors of the zebras, which are now confined to

equatorial Africa, extended to the extreme north of the continent. To

the same period belongs a wild ass very similar to the Ethiopian ass

(#. asinus), an animal which survived in this region until exterminated

by the Greeks and Romans, and is now confined to the highlands of

Abyssinia.

Among the Artiodactyla the presence of camels’? (C. thomasi) in

Paleolithic Pleistocene times and even in deposits of Neolithic age (C.

dromedarius) is extraordinarily interesting. There is no evidence as to

domestication. ‘The earlier of these two camels of ancient Libya had

longer legs and was of heavier build than the dromedary. ‘The rare re-

mains of the later form, probably identical with the recent dromedary,

may be those of a race which was already emigrating or becoming extinct.

The presence of the camel is one of the most convincing proofs of con-

nection of this fauna with that of the Upper Siwaliks of southern Asia,

and thus of North America.

Especially significant of Asiatic and Siwalik affinity are the Pleisto-

cene cattle and buffaloes of north Africa, including contemporary species

of Bos, all belonging to late Quaternary or to the Neolithic age, partly

domesticated, and with remote resemblances to the Pleistocene cattle of

France and Spain. Similar Asiatic affinity is found in the remains of a

buffalo (Bubalus antiquus) allied to the existing Indian form; this was

a powerful beast which presumably lived in herds, frequenting grassy

plains and swampy districts, and in its presence here we seem to find con-

firmation of what geology teaches us in regard to the dampness of the

Quaternary climate. The disappearance of the buffalo from north Africa

at the commencement of the Recent Period was no doubt due to the in-

creasingly dry conditions, and partly to destruction by man.

The great number and variety of antelopes is most astonishing in this

region, which at present is inhabited only by the gazelles (Gazella), the

hartebeests (Bubalis) and addax antelopes (Addaxr). The Pleistocene

fauna includes gnus (Connochetes), several species of Bubalis still rep-

resented in the Barbary States, an aberrant form (Oreonagor), related

to the nilgai of India, nine species of gazelles (Gazella), the oryx

(Ory), the nabor (Cervicapra redunca), several large elands (Oreas),

such as now inhabit south Africa, as well as dwarf antelopes (Cephal-

ophus). Beside these plains and desert types of ruminants, the hills

were covered with wild sheep (Ovis palwotragus) very similar to the ex-

isting Barbary sheep, as well as goats (Ovis promaza).

4 PomMEL, A.: “Caméliens et Cervidés,’’ Carte Géol. Algérie, Paléont. Monogr. Algiers,

1893.

“e

28 ANNALS NEW YORK ACADEMY OF SCIENCES

row)

In the rivers there lived in early and later Pleistocene times a series

of species of hippopotami (fH. hipponensis, H. sirensis, H. icosiensis)

leading to a form (H. annectens) related to the existing Nile hippopota-

mus. There are also two types of wild boar (Sus), and more abundant

than these were the wart-hogs (Phacocherus) found in the caves and

alluvial deposits of Barbary.

Preying upon these Herbivora were lions, leopards and hyzenas, which

are compared by Pomel with Pleistocene cave forms of Europe. There

are also jackals, wolves, the ichneumon and, possibly, a polecat.

Fic. 2.—Skeleton of the Pleistocene pigmy hippopotamus of Madagascar, Hippopotamus

madagascariensis, together with a skull of the recent hippopotamus, H. amphibius

In the American Museum of Natural History.

African-Huropean Distribution.—Of this imposing list the following

types occur both in Africa and in the Lower and Middle Pleistocene of

Europe, the species being similar if not in some instances identical.

Southern elephant (#. meridionalis), which is also found in Pliocene and early

Pleistocene deposits of Europe.

HWlephants similar to HL. antiquus of Europe and its dwarf representatives in

Malta and other Mediterranean islands are found in the Upper Pleistocene

deposits of north Africa.

Long-headed rhinoceroses. It would appear probable that the woolly rhi-

noceros (D. antiquitatis) which is closely related to the ‘‘white”’ rhinoceros

(D. simus) originated in Africa, but no animal resembling it has been

discovered in the African Pleistocene.

OSBORN, REVIEW OF THE PLEISTOCENE 229

One of the Pleistocene horses of north Africa (2. numidicus) is closely similar

to the Upper Pliocene EH. stenonis of Europe.

With these animals may have come the lion (Felis leo) which was widely

spread over southern Europe..

Hynas. The striped (H. striata) and spotted hyznas (H. crocuta) are com-

mon to Europe.

Bears. The bear (Ursus lybicus) found fossil in Algeria seems to belong to

the Helarctos group, possibly derived from the small U. etruscus of the

European Pliocene and now represented by the Malayan sun bear.

Macherodonts. Recently (Stromer) sabre-tooth tigers have been discovered

in Pieistocene Egypt.

Primates. The primates are represented by the macaque (Macacus), not very

different from the existing forms which frequent the region of the Straits

of Gibraltar. In Pleistocene times the macaques ranged northward into

southern France (Harlé).

Suillines. Wild boar (Sus) may have affinities with the Pliocene types of

Europe.

With the exception of the above list, there is little in common between

the large fauna of north Africa and that of Europe in Pleistocene times.

AFRICA IN PALXZOLITHIC AND NEOLITHIC TIMES.

Giraffes very similar to the recent African giraffe (C. giraffa) have

been found in mid-Pleistocene deposits associated with Paleolithic stone

implements of the Chellean type. Industry'® similar to the Chellean

but not necessarily of the same age is found in Africa from Egypt to the

Cape. Giraffes are also depicted in rock drawings of Neolithic age in

Algeria.

In Neolithic times there existed at least one species of deer, whereas at

present there are two kinds of deer, the red and the fallow,** in north

Africa, both undoubted Eurasiatic migrants.

The prehistoric men of the Barbary States apparently obtained and

domesticated the horse, species of sheep and several dogs, and left many

sketches of animals on the rocks of the region.*®

PLEISTOCENE OR GLACIAL EPpocH IN EUROPE

After the establishment of the single glacial theory by Charpentier and |

Louis Agassiz (1836-1840), there gradually developed in Europe and

122 PaLLARY, P.: “Note sur la Girafe et le Chameau du Quaternaire Algérien,”’ Bull.

Soc. Géol. France, Ser. 3, Vol. XXVIII, pp. 908-909, 1900.

18 OBERMAIER, HuGo: *“‘Der Mensch der Vorzeit. Band I. of Der Mensch aller Zeiten,’

Alleg, Verlags-Gesellsch. m. b. h. Berlin, Munich, Vienna, 1912.

14 See LYDEKKrR, RICHARD: Deer of all Lands. The North African red deer (Cervus

elaphus barbarus) is smaller than the European race. Evidence on the range of the com-

mon fallow deer (Cervus dama) in northwestern Africa is not very full.

15 See PoMBL, '93, ’94, '95, ’96, ’97, ’98.

30) ANNALS NEW YORK ACADEMY OF. SCIENCES

Cys

America the hypothesis of several glacial advances of varying duration

and severity alternating with interglacial temperate periods during which

the ice retreated and conditions of climate prevailed which in some in-

stances were even milder than the present in the same latitudes.

As early as 1856 Morlot observed a relatively warm flora between two

Swiss glacial deposits at Diirnten, and he subsequently advanced a theory

of three glacial stages. James Geikie (1871-1894) developed the hypoth-

esis of a succession of six glacial and five interglacial stages and climates.

In 1883 Boule from his observations along the Mediterranean coast main-

Young drift Limit——-—=

Middle Dnft Umit ———

Old Drift Limit ++++¢4

| Scale 1:5,000.000

joo so conn |

Fic. 3.—Glacial map of northern Germany and the Netherlands

This map shows the drift and terminal moraines of Glaciations I-II, Scanian + Saz-

onian (old drift), III Polandian = Riss (middle drift), IV Mecklenburgian = Wiirm.

(upper drift). After Leverett, 1910.

tained that there is evidence of three great glacial advances, the first fall-

ing within the close of the Pliocene Epoch, the second falling properly

_ within the Pleistocene. The firm foundation of the quadruple theory in

Europe was laid, however, by the researches of Penck and Briickner** in

the Alpine region, published in 1909. According to this classic work the

entire Glacial Epoch is assigned to the Pleistocene or Quaternary Period.

Its deposits include the entire “Diluvium” and “Drift” of earlier geolo-

gists.

All the river gravels, boulder-clays and moraines of the Glacial Epoch

1€ PENCK, ALBRECHT, and BRUCKNER, EpovarD: Die Alpen im BEiszeitalter. III. Die

BHiszeiten in den Sudalpen und im Bereich der Ostabdachung der Alpen. Leipzig, 1909.

OSBORN, REVIEW OF THE PLEISTOCENE " 934

are of later date than the marine Pliocene deposits of southern Europe.

Before any of these glacial deposits occurred there was an elevation of

the marine Pliocene strata along the southern Alpine borders from sea

level to a height of from 300-500m.; there also occurred erosion of these

marine strata by rivers. Thus in the valley of the Po there is a consid-

erable time interval between the closing marine conditions of the Plio-

cene and the opening Pleistocene conditions. In the valley of the Rhone

also there is a marked interruption between the strata of the Pliocene and

of the Glacial epochs, the latter overlying the strata recognized as Upper

Pliocene, which in turn overlie the marine Pliocene.*” }

This interval between the Pliocene and Pleistocene corresponds with

very important changes which occur in the mammalian life of Europe,

namely, in the extinction of many characteristic Pliocene mammals, such

as the anthropoid apes, the antelopes and the mastodons.

a Sierra de Gredos Alps Mis

nwo, [3 3 Atlas Mts ee Mts. |

secel [al > | France Germany Scandinavian Plateau

ee i= |= | | Denmark

— = +

| | North Cape

Strait of Gibraltar Garonne hone North Skeger

- Valley Valley Sea Fak

PROFILE OF PAST AND PRESENT SNOWLINES AND CLACIERS OF EUROPE

Fic. 4.—Theoretic snow levels during the Glacial Epoch

Prepared under the direction of the author by Chester A. Reeds from data given by

J. Geikie, Penck, Briichner, Leverett and Stieler’s Hand-Atlas, December, 1914.

The traces of four different glacial advances and retreats observed

around the northern slopes of the Alps by Penck and Briickner*® were fol-

lowed with their “river drifts” and moraines down the Danube to the

neighborhood of Vienna; they were found to be clearly marked in the re-

gion of the upper Rhine and of the Rhone around Lyons, and distin-

guishable both by the greater or the less extension of their borders and by

the greater or less erosion which has occurred in the intervals between

their successive depositions. These four advances were named respec-

tively the Giinz, the Mindel, the Riss and the Wiirm.

As an instance of the disparity between the duration of these several

glacial advances with the accompanying descent of the ice and snow line,

the old moraines of the Riss or third glaciation form a girdle around the

more recent Wiirm or fourth glaciation, proving that the Riss was not

7 Op. cit., pp. 654-655.

1480p. cit., p. 47.

232 ANNALS NEW YORK ACADEMY OF SCIENCES

only a more extensive glaciation but that the snow line was 100m. lower.

It is also estimated that the climate of the Riss was one-twelfth more

severe than that of the Wurm.

In northern Germany only three great glacial advances are recorded,

while still farther north, in Scandinavia, there was in a sense only one

Glacial epoch, since the ice cap never retreated so far as to permit of

interglacial deposits. This is in accordance with the anthropological

fact that only toward the close of Postglacial times does Scandinavia

show traces of human habitation in the arrival of the Neolithic men;

whereas in France and Germany there is evidence of human habitation

as early as the Second and Third Interglacial Stages.

In the meantime American geologists have also discovered similar

proofs of four successive glacial advances and more temperate inter-

glacial stages. The correlation of these conditions in the New and Old

Worlds suggested by Penck, Chamberlin and others has recently been

reviewed with great precision by Leverett,1® to whose work we shall fre-

quently refer. The most recent results of geologic and anthropologic

correlation with some original modifications are graphically presented in

the accompanying diagram (Fig. 5) by the author and Reeds.”°

The river terraces are of great importance both in geology and in an-

thropology. In general the “high terraces” belong to the earlier glacia-

tions and the “low terraces” to the latest. Thus the “high terraces” of

the Alpine region belong to the Riss or glaciation III; in the valley of

the Rhine the “high terrace” is visible near Basle; the “low terrace” of

the Wiirm or glaciation IV occupies vast surfaces on the upper Rhine

and contains a mammoth (F£. primigenius) fauna. The “high terraces”

in the Paris basin reach 30m. above the level of the Seine, while the “low

terraces” are only 5m. above the level of the Seine and subject to floods ;

the “high terraces” in the valley of the Seine contain the First Inter-

glacial (EZ. meridionalis, EZ. stenonsis) fauna, while the “low terraces” of

the Seine and of the Somme contain the Second and Third Interglacial

fauna (EF. trogontheru, EL. antiquus, and D. merckit).

DURATION OF THE PLEISTOCENE

The Pleistocene was estimated by the American geologist Dana (1874)

_to be equal to about one-fourth of the entire Cenozoic Era, 1. e., 700,000

years. By Ward (1885) and Williams (1895) it has been estimated at

19 LEVERETT, FRANK: “Comparison of North American and European Glacial Deposits.”

Zeitschr. f. Gletscherkunde, Vol. iv, pp. 241-316. 1910.

°° REEDS, CHESTER A.: Dr. Reeds has prepared the climatic curve from data furnished

by Penck, Leverett, Taylor, Chamberlin, Salisbury, Geikie, Schmidt, Coleman and Osborn.

Dated October, 1914.

OSBORN, REVIEW OF THE PLEISTOCENE 233

|CORRELATION Of me RACIAL, CULTURE & LIFE STAGES 19/4

PR ETL A

POST GLACIAL ZZDAU Ss VEOLIT) te RECENT FOREST, MEADOW, ALPINE

"Newer Loess’ ZGSCHNIT.

WETLE | eee eee

SLLLLELEALLLABOAL

GRO'MAGNON | PeINDEER PERIOD, ARCTIC

IV. GLACIAL 7-22

WURM, WISCONSIN” ZLAUA

WEANDERTHAL| TUNDRA, STEPPE , ALPINE

” " FOREST, MEADOW

“Upper Drift” ZZ ;

“Lowest ferraces ae Zs COLD FAUNA

ARRIVAL: STEPPE , TUNDRA, FAUNA

“ (KRAPINA)| (AST WARM AFRICAN-ASIATIC

3. INTER- Ai ae |

GLACIAL 7% LITHIC E.ANTIQUUS, HIPPOPOTAMUS |

RISS -WURM 4 ijt D.MERCKII, E.TROGONTHERII |

SANGAMON titi A ALSO FOREST, MEADOW |

pee Loess” EURASIATIC FAUNA

WEGLACIAL %

COLD TUNDRA FAUNA

Mit - 2 oe al WOOLLY MAMMOTH &

As RHINOCEROS. FIRST

STEPPE & REINDEER

ae ae a Z a f.

TLLINOIAN FY Ai

Ze ee ees

2. INTER- tt aR care oT

aoe i WARM AFRICAN-ASIATIC |

HELVETIAN ini

YARMOUTH ‘ E.ANTIQUUS , E. TROGONTH-

Long Warm i! HEIDELBERG) eR 6. MERCKII, HIPPO-

ti;

Stage Me | 171275000 POTAMUS

‘Older Loess’ zt

ates 1 A 4

RaN

tj BAN

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MINDEL , SAXONIAN

KANSAN

‘Old Drift’

Mii tiiiié 123 yy, ee

I INTER- ZY ae ae te

GLACIAL

NORFOLKIAN | ij

GUNZ-MINDEL

AFRICAN - AS/IAT/C FAUNA

£. MER/D/IONALIS —TROGON -

THER//, D. ETRUSCUS,

HIPPOPOTAMUS

Hyp. ae Pe ee

LGLacia & 3

Z COLD FOREST BED

IAN FAUNA /N S. BRITAIN Pt Megha

"Old Terraces’ isis See nia

EEL, ld SE a rs ics PLIOCENE

PLIOCENE Zl [ WARM FOREST

GLAC/AL

AND

Fig. 5.—Divisions and contemporaneous events of the Glacial Epoch

Prepared under the direction of the author by Chester A. Reeds from data observed

and correlated by J. Geikie, Penck, Chamberlin, Salisbury, Leverett, Schmidt, Coleman

and Osborn. October, 1914.

W YORK ACADEMY OF SCIENCES

)

ANNALS NE

234

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OSBORN, REVIEW OF THE PLEISTOCENE 235

cne-third the entire Cenozoic, 1. e., 1,000,000 years. If with Wallace we

accept Croll’s theory and estimate, the last glacial advance would date

back to the last period of great eccentricity of the earth’s orbit, namely,

200,000 years, but this we now consider excessive. The following figures

show the variations of opinion on this subject and the two opposite tend-

encies of greatly expanded or greatly abbreviated estimates of Pleistocene

time:

Lyell, “Antiquity of Man’”..... 1863 800,000 years

NOG INRA sain. hc eantnct Re ea ice cere UY 1893 100,000 *

RMT PL OOM oe osiirs <retoceteyaalemereers sa so's =o 1893 400,000 ‘*

STL ISSR Pe Shana Ca eines ara 1900 400,000 <

1 ESTING 0 aoe SP Poa hehe a i ta oa 1909 520,000 to 840,000 years

The very high estimate of 840,000 years made by an eminent and

usually conservative authority such as Penck appears excessive unless we

are to expand our estimates of Tertiary time (see p. 63) to 20,000,000

years and of the pre-Tertiary into hundreds of millions of years.

All the arguments for the briefer estimates of Pleistocene time have

recently been brought together by Wright.’

Antiquity of Man.—Vast interest attaches to this duration problem in

connection with the antiquity of man. In the calculations of Penck?? the

time since the Fourth or Wiirm glaciation has been used as a measure-

unit to calculate the length of the previous glacial and interglacial

periods. It is believed that since the climax of the Wiirm glaciation

from 20,000 to 34,000 years have elapsed. Geologic, prehistoric and

historic events since the close of the last glaciation make this estimate

appear not excessive. In regard to the previous time intervals, the au-

thor does not pretend to give an absolute age estimate, but simply a

survey of the relative magnitude of the time periods with which we are

dealing.

The unit of measurement is Postglacial or post-Wiirm time which

Penck**® estimates at 20,000 years. On the basis of this estimate the

time (520,000 years) covered by the whole Glacial Epoch is relatively

distributed as follows:

21 WRIGHT, G. FREDERICK: The Ice Age in North America and its Bearings upon the

Antiquity of Man. S8yo. Bibliotheca Sacra Co. Oberlin, 1911.

2 PENCK, A.: “Das Alter des Menschengeschlechts.” Zeitschr. Ethnol., No. 3, pp:

390-407. 1908.

3 PENCK, A., and BRUCKNER. Epovarp: “Die Alpen im WHiszeitalter. Dritter Band. Die

Hiszeiten in den Sudalpen und im Bereich der Ostabdachung der Alpen.” S8vo. Tauch-

nitz, Leipzig, 1909. (pp. 1153-1176, ““Chronologie d. Eiszeitalters in d. Alpen.” Penck.)

36 ANNALS NEW YORK ACADEMY OF SCIENCES

duration Totals, ayer

Units Years Years Meters

Postglacial, post-Wiirm hemicycle... 1 20,000 20,000 aia

LV. (0f | WERMAGIACIS TION Uo ok ye oe 1 20,000 40,000 1,200

3rd or Riss-Wiirm Interglacial Stage. 3 60,000 100,000 Sane

TH -or Herss:GrxGra tion J]. aces Seas 1 20.000 120,000 1,250

2nd or Mindel-Riss Interglacial Stage 12 240.000 360,000 agers

i or: MinpEEn GG EACrATION: ooo See ce eee eee il 20.000 380,000 1,306

1st or Giinz-Mindel Interglacial Stage 5 100.000 480,000 aged

for. ‘Gunz \Giacrenion «ih fv eee eee i 20,000 520,000 1.200

The three chief conclusions of Penck are as follows:

1. If the whole Ice Age extended over a period of 500,000 to 1,000,000

years, the Second very long warm Interglacial Stage, also known as the

Mindel-Riss or Helvetian, is reckoned at more than 200,000 years, while

the final relatively short interglacial stage, the Riss-Wiirm, is reckoned

at 60,000—100,000 years.

2. The duration of the Lower Paleolithic culture periods, the pre-

Chellean, Chellean and Mousterian, would by this reckoning be much

longer than that of the Upper Paleolithic culture periods, the Aurig-

nacian, Solutrean and Magdalenian. Penck estimates that since the be-

ginning of Magdalenian times 24,000 years may have elapsed and since

its close about 16,000 years.

3. Compared with these prolonged Paleolithic divisions the Neolithic

Stone and Metal periods have occupied an almost unappreciable length

of time. If the beginning of the Neolithic lake dwellings is dated about

5,000 to 7,000 years ago we estimate that the beginning of the Copper

Age in Europe dates back between 3,000 and 3,500 years; in Africa it is

much more ancient.

The human culture stages are arranged above not according to Penck

but according to the more recent correlations of Obermaier, Breuil,

Schmidt and others.

GEOLOGIC AGE OF THE CULTURE STAGES

The trend of Paleolithic research lately has been to draw all the

human culture periods from the pre-Chellean to the Magdalenian closer

together and to reduce the time assigned for their evolution. All the

French authorities, led by Boule, Cartailhac, Breuil and Obermaier, are

now agreed in assigning the earlier Palwolithic cultures, the pre-Chel-

lean, Chellean and Acheulean, to the Third Interglacial Stage and not to

the Second. Schmidt has also lately declared himself in favor of this

view after a most exhaustive and valuable investigation of this problem.

OSBORN, REVIEW OF THE PLEISTOCENE 937

A summary of the correlation presented in this chapter is embodied in

the Table of Osborn and Reeds above. A summary of the very diverse

opinions on this subject is embodied in the Table of Wiegers below.

A very strong reason for abbreviating our estimates of the period

which has elapsed since the appearance of man of the pre-Chellean ecul-

ture stage in Kurope is found in the relatively unchanged condition of

the river valleys of the Somme in northern France and of the Vézére in

Dordogne, in which the earliest human cultures occur. ‘The Vézére has

not materially changed since Acheulean times. The pre-Chellean, Chel-

lean and Acheulean specimens found in the Somme valley are also con-

nected with the present river system. Both on the Somme and the

Marne the Chellean and pre-Chellean cultures occur on the “lowest ter-

races.” Again, there is no faunal break between late Chellean and early

Acheulean times nor between late Acheulean and early Mousterian times.

The first great faunal break is that produced by the Fourth glaciation.

In favor of Penck’s contention as to the earlier geologic age of the

Chellean is the occurrence of pre-Chellean and Chellean paleoliths in

association with a very primitive mammalian fauna such as is character-

istic of Second Interglacial times.

ELEVATION AND SUBSIDENCE OF LAND IN QUATERNARY TIMES

The relations of the mammals of Europe with those of Asia on the east

and Africa on the south were profoundly affected in Pleistocene times by

the periods of elevation of the continental shelf, resulting in the creation

of new land connections which facilitated migration, or of subsidence

which cut off and isolated many migrating forms from their centres of

origin and dispersal. The maximum elevation, as represented in the

accompanying diagram (Fig. 1), never occurred in all portions of the

continent of Hurope at the same time, because there were oscillations

both on the northern and southern borders of Europe and Asia.

The beginning of the Pleistocene Epoch is one of elevation and is re-

markable for the broad land connections between Europe, Africa and

Asia. It represents the last stage in that vast community of mam-

malian life which during Pliocene times distinguished the entire region

of Europe, Asia and Africa.

The theoretical relation which elevation and subsidence respectively

bear to the glacial and interglacial stages and phenomena is, broadly

speaking, as follows:

ELEVATION, emergence of the land from the sea, broad land connections

facilitating migration, retreat of the ice caps. periods of erosion of the

river valleys and formation of terraces.

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OSBORN, REVIEW OF THE PLEISTOCENE Dag

SUBSIDENCE, submergence of the land and advance of the sea, land con-

nections interrupted, advance of the glaciers, periods of deposition and

filling of the valleys.

Thus Boule and Geikie consider the glacial stages as mainly periods of

continental subsidence and filling of the valleys, the interglacial as times

of elevation and erosion of valleys and terraces. Penck estimates the

elevation of southern Europe at 100m. in the beginning of Pleistocene

times ; he speaks of the elevation of the Alps during the Second Inter-

glacial Stage. Geikie describes the southern half of the North Sea as

dry land during the First Interglacial Stage traversed by a northern ex-

tension of the River Rhine, while the approach of the Second Glacial

Stage was heralded by a submergence of this area of the North Sea.

Again (Geikie) during the Second Interglacial Stage the English Chan-

nel and probably a large part of the North Sea became dry land. Finally

(Pohhg) during the Third Interglacial Stage there was a period of

continental elevation and a dry, cold steppe climate in western Europe.

Consistent with this hypothesis is the deposition of loess during the

Second and Third Interglacial Stages, also during the Postglacial Stage,

because loess deposition is characteristic of dry and elevated continental

climates with winds prevailing in one direction.

Final Subsidence.—Consistent with this hypothesis also is the fact that

general and local swbhsidence in the northern hemisphere was the chief

feature of closing Pleistocene times or the very cold Postglacial Stage;

all the old continental connections which had been characteristic of the

Tertiary were cut off; in the northwest the English Channel was formed,

Great Britain became isolated from Europe, Ireland first lost its land

connection with Wales and then with Scotland ; to the eastward the Medi-

terranean Sea extended into the A¥gean region and cut off the old land

connection between Greece and Asia, which had so long served to connect

Greece with the mammalian life of southern Asia. During a period of

extreme subsidence, the Black Sea, the Caspian, the Sea of Aral formed

a large single sheet of water known as the Hyrcanian Sea. In southern

Asia similar subsidence and separation phenomena were in progress; the

islands of the East Indies, Sumatra and Java were cut off from the

Malay Peninsula. The separation of the Japan and Philippine archi-

pelagos probably occurred in late Postglacial times. To the far north-

east late in Pleistocene times Asia lost its connection with America,

Bering Straits were reopened, and the so-called Nearctic region of North

America was completely isolated from the Palearctic region of Eurasia

after a long period of community and free intermigration of Holarctic

~ life.

240 ANNALS NEW YORK ACADEMY OF SCIENCES

Final Elevation—Vhis extreme subsidence was followed in late Post-

glacial times both in Europe and America’ by reélevation which gave the

continents their present contours and climates.

ALTERNATIONS OF CLIMATE AND FLORA

Fluctuations of temperature and of moisture in Pleistocene times are

indicated first by the advances and retreats of the ice caps, second by the

presence of arctic or temperate Mollusca in the coast waters, third by the

variations in the flora in glacial and interglacial times, fourth by the

alternate appearance of the northerly or southerly types of mammals and

birds, fifth by the nature of the geologic depositions, sixth by the nature

of the land Mollusca in the loess. Combining the evidence derived from

these various sources the theoretic broad divisions of the climatic se-

quence are as follows: (1) the cold and moist phases connected with the

successive glacial advances and retreats; (2) the warm to temperate

climates of the First and Second Interglacial Stages and first half of the

Third Interglacial; (3) the dry and cold climate of the second half of

the Third Interglacial Stage and early Postglacial times; (4) the damp

and cool climate of late Postglacial times favorable to forests.

The theoretic alternating conditions of each complete glacial cycle are

as follows :?4

Subsidence: Glacial Maximum: Tundra flora and fauna

Glacial Retreat: Cool and moist forest flora and fauna

Elevation : Interglacial : Dry conditions, flora and fauna. Steppe

Glacial Advance: Cool forest fauna and flora

Subsidence: Glacial Maximum: Cold tundra flora and fauna

Low Glacial Stage Temperatures.——Low temperatures during the pe-

riods of glacial advance are attested both by the advent of northern Mol-

inusca, marine and terrestrial, northern flora, and the repeated arrival in

Europe of members of the cold fauna of the arctic tundras, including both

the smaller and the larger mammals and the birds, as well as the cold

fauna of the high, arid steppes of western Asia. Low temperatures are

attested also in early Postglacial times during the great Aurignacian-

Magdalenian art period by the heavy covering of hair indicated on all

the animals depicted by the Upper Paleolithic artists. This hairy cov-

ering coincides exactly with that of the extreme northern tundra types of

reindeer, woolly rhinoceros (D. antiquitatis) and woolly mammoth (2.

primigentus) found imbedded in the ice or frozen soil.

% Compare Wiist.

OSBORN, REVIEW OF THE PLEISTOCENE 941

Moderate Estimates of Temperature-——That the advancing glaciers

alone do not constitute proof of very low temperatures is observed in

Alaska, where very heavy snowfall or precipitation causes the accumula-

tion of great glaciers, although the mean annual temperature of the

glacier region is 40°-45° F. (4.44°-7.22° C.) as compared with that of

northern Germany, 45°-50° F. (7.22°-10° C.), « ¢., from mouth of

Rhine S. E. along source of northward flowing rivers, e. g., Elbe, Vistula,

etc. Neumayr estimated that during the Ice Age there was a general

lowering of temperature in Europe of not more than 6° C., and held that

even during the glacial advances a comparatively mild climate prevailed

in Great Britain. Martins estimated that a lowering of temperature to

the extent of 4° C. would bring the glaciers of Chamonix down to the

level of the plain of Geneva. Penck estimates that all other atmospheric

conditions remaining the same as now a fall of temperature to the extent

of 4° to 5° C. would be sufficient to bring back the Glacial Epoch in

Hurope. Perhaps the strongest proof that Europe was not refrigerated

during the first, second and third glacial advances is the survival of the

African-Asiatic fauna throughout the whole period until the fourth

_ glaciation, which was accompanied by widespread severity of climate.

Warm and Temperate Interglacial Stages—Similarly the early hy-

potheses of extremely warm or subtropical conditions, based chiefly upon

the northerly distribution of hippopotami and rhinoceroses, animals

which we now associate with tropical conditions, are not supported by the

study of the interglacial flora. It is quite probable that both the hippo-

potami and rhinoceroses of the so-called “warm, fauna” were covered with

hair although by no means so thickly covered as the woolly rhinoceros

and elephant of the arctic tundras. There is evidence that in the First

Interglacial Stage southern England and France enjoyed somewhat

warmer and moister conditions of climate than the present. The Second

Interglacial Stage also, which is commonly distinguished as the “long

warm” Interglacial Stage, was of somewhat higher temperature than the

present. ‘The general evidence is that both in Europe and North Amer-

ica, especially in the First and Second Interglacial intervals, the climate

in the northern hemisphere was somewhat more equable and milder than

at present, with a higher mean temperature, at certain intervals with

greater precipitation of moisture, at other intervals much more cool and

arid. There was perhaps more sunshine than now.

As a result of favorable interglacial conditions arboreal vegetation

flourished to the far north along the Arctic ocean, and the present tundra

regions of Siberia and British America then supported forests which

949 ANNALS NEW YORK ACADEMY OF SCIENCES

have long since been extirpated, the northern hmit of similar living trees

now lying far to the south.

Alternations of Flora.—lt is clear from these great successive fluctua-

tions of temperature, moisture and aridity during Pleistocene times that

the flora cannot be treated as a unit nor as progressing in a single direc-

tion like the flora of preceding epochs; the flora as well as the fauna

presented alternations of arctic, boreal and temperate species which

migrated southward and northward following the advances or retreats of |

the glacial cap. Thus we may observe evidences of changes of climate

and flora from forested conditions to steppe conditions and back to for-

ested conditions. From the beginning of the Fourth Interglacial in-

terval to the present time, the Alps region (Penck, Brickner, 1909) has

apparently gone through a cycle of changes such as the following:

VEGETATION CLIMATE EpocH

Fourth, forest conditions Western European, oceanic Modern

Third, steppe conditions Western Asiatic, continental Fourth Glacial and

Postglacial

Second, tundra conditions Northeastern-European, Fourth Glacial

subarctic

First, forest conditions Western-European, oceanic Third Interglacial

The elephants (Hilzheimer, 1913) in the structure of their grinding

teeth afford clear indications of the plant life, whether consisting mainly

of grasses or forests, but not of climate except in so far as vegetation is

dependent upon moisture. ‘The advance and retreat of the ice is de-

pendent both upon moisture and extreme cold and involves the frozen

subsoil conditions of the tundras which are fatal to forests. Cases of

alternation of conditions favorable to Hlephas trogontheru, which is be-

heved to be a grass-eating form, and of Hlephas antiquus, which is be-

heved to be a forest-living form, are observed in Taubach by Wiist.*°

This author observes in the lower layers Hlephas antiquus succeeded in

the middle layers by H. trogontherii and then in the upper layers again

by H. antiquus, and deduces from, this succession a change of conditions

from forest, to steppe, to forest.

FAUNAL LIFE ZONES OF EUROPE

In the whole history of the Mammalia in various parts of the world we

know of no conditions so unusual and complex as those which prevailed

in Hurope in Pleistocene times. These conditions were the product of

> WUST, EWALD: “Die plistozinen Ablagerungen des Travertingebietes der Gegend von

Weimar und ihre Fossilienbestiinde in ihrer Bedeutung fiir die Beurteilung der Klimasch-

wankungen des Hiszeitalters.’”” Zeitschr. Naturw., Bd. 82. pp. 161-252. Leipzig, 1911.

OSBORN, REVIEW OF THE PLEISTOCENE — 9AZ

cycles of environment and of life which have never prevailed before and

will never recur even if the world were to enter a fifth glacial stage, for

besides the extraordinary geographic and climatic changes which have

been outlined in the previous pages there was the prodigal profusion of

life which survived from Pliocene times and has since become extinct.

The result of these complex conditions was the assemblage in Europe

of animals indigenous to every continent on the globe except South

: a |

rae

li ald

ny !

17 f

n rm

inh

eee Ayes

see

Pic, 6.—Five chief zoédgeographic regions of Europe, Asia and northern Africa from which

the mammals migrated into western Europe during the Pleistocene

America and Australia, and adapted to every climatic life-zone from the

warm and dry plains of southern Asia and northern Africa to the tem-

perate forests and meadows of Eurasia, from the alpine heights of the

Alps, Pyrenees and Altai Mountains to the high, dry steppes of central

Asia with their alternating heat of summer and cold of winter, from the

tundras or barren grounds of Scandinavia, northern Europe and Siberia

to the mild climate of southern Europe. All these animals had been

evolving during the Pliocene Epoch in these various habitats and they

QAA ANNALS NEW YORK ACADEMY OF SCIENCES

also underwent a very considerable evolution during Pleistocene times in

their respective habitats.

-Owing to successive migrations and invasions into Europe of these

exotic types of the north and south we should not expect to find a con-

tinuous phyletic evolution or transformation such as we have observed in

the earlier epochs, excepting only that which occurred among the Eura-

siatic forest and meadow types which appear to have been native or in-

digenous in Europe from the earliest Pleistocene to prehistoric times.

These Eurasiatic forest and meadow mammals were continuous residents,

retreating in the coldest periods to the shelters on the east and south.

Cervus elaphus, for example, passed through several subspecifie stages of

evolution. The invading hordes from the tundras, the steppes, from

northern Africa and from Asia represent branches which had their evolu-

tion elsewhere. This is true both of the mammals and of the races of

men which had their genesis in the far east and southeast ayd arrived in

Europe when it was a fertile peninsula, a region too small to be the seat

of a continental evolution or adaptive radiation.

The five great regions which contributed to the European Pleistocene

were as follows:

AFRICAN AND ASIATIC, PLAINS AND FoREST TYPES

EUROPEAN AND ASIATIC, FOREST AND MEADOW TYPES

EUROPEAN AND ASTATIC, ALPINE TYPES

STEPPE REGIONS OF SOUTHERN EURASIA AND EASTERN SIBERIA

Arctic TUNDRA REGIONS OF NCRTHERN EURASIA

Interature-——The African-Asiatic element in these Pleistocene faunas

was the first to be recognized and commented upon by the early writers;

it is commonly known as the “warm fauna.” We owe especially to Neh-

ring the discrimination between the tundra and the steppe faunas.

Gaudry, Harlé, Woldrich, Studer and Boule have added to our knowl-

edge of these faunz. Other contributors are Pohlig, Soergel, Forster,

Hilzheimer, Wiist and Dietrich. A strict systematic revision and intro-

duction of the trinomial system is greatly needed. The most complete

recent faunal lists of the late Pleistocene deposits in which traces of man

are found are those of Koken and Schmidt (1912), who have also insti-

tuted the closest correlation between the migrations of the Mammalia

and the successive stages of human culture.

African—A siatic Mammals, Warm Fauna.—These mammals inelude

those which first appear in the Upper Pliocene and survive into Lower

Pleistocene times; also those which first appear in the Second Inter-

glacial Stage and constitute the so-called “warm fauna” which survived

in Europe until the middle or close of the Third Interglacial Stage. The

OSBORN, REVIEW OF THE PLEISTOCENE 945

principal members of this list together with the probable continental cen-

tres of their origin are as follows:

Macacus ? sp. (Africa)

Elephas meridionalis (Asia)

Elephas antiquus (Asia-

Africa)

Dicerorhinus etruscus (Asia)

Pe merckii (Asia )

Hippopotamus major (Asia)

Bos primigenius (Asia)

Bison priscus (Asia)

Equus stenonis

Macherodus latidens

Hyena spelea ( Asia-Africa )

“striata (Asia-Africa )

Cams aureus (Africa)

Felis leo antiqua (Africa)

“ pardus

Sudelefant

Aitelefant

Nashorn

Mercksches Nashorn

Flusspferd

Urochs, Auerochs

Wisent, “‘Auerochs”

Hohlenhyane

Schakal

Altl6we

Leoparde

Macaque baboon

Southern mammoth

Straight-tusked elephant

Etruscan rhinoceros

Broad-nosed rhinoceros

Hippopotamus

Urus

Primitive bison

Upper Pliocene horse

Sabre-toothed tiger

Cave hyzena

Jackal

Lion

Leopard

The remains of these animals play a very important part both in the

subdivision of the geologic horizons of Hurope and in theories regarding

the alternation of climates, as well as in the determination of the an-

tiquity of man. They are found chiefly in the river sands, river deposits

and “High Terraces” of the First and Second glaciations and “Low Ter-

races” of the Third Glacial and Interglacial Stages. Some survivors are

found in the shelter and cavern deposits of the Third Interglacial Stage

as objects of the chase. Only two of these animals, the urus and the

bison, survive to become members of the Prehistoric Forest and Meadow

Fauna of Europe. The lion also survived into Postglacial times but dis-

appeared in Europe before the Prehistoric and Neolithic periods.

Although originally derived from southern Asia or from Africa, the

woolly elephant (HL. primigenius) and woolly rhinoceros (Diceros antiq-

uitatis) are not to be placed with the African-Asiatic fauna because they

appear in Europe only with the northern Glacial or Tundra Fauna and

are invariably indicative of cold climatic periods.

Eurasiatic Forest and Meadow, Temperate Fauna.—These animals were

resident in the forests and meadows of Europe during the entire Pleisto-

cene Epoch and survived with a few exceptions into Postglacial and Pre-

historic times. In fact, they are probably separable by differences of

specific and subspecific value from their successors in prehistoric Europe,

but many authors still embrace them within the existing specific terms.

They are divided into Forest and Meadow types, the former with brachy-

odont teeth adapted to browsing habits and forest environment, the latter

with elongate or hypsodont grinding teeth adapted to meadows and a diet

246 ANNALS NEW YORK ACADEMY OF SCIENCES

of grasses. Some of them, like the beavers, are stream- and river-dwellers.

Naturally there is not always a sharp line of division between the habitats

of these Forest and Meadow types: the Carnivora especially wander after

their prey from the forests into the meadows and along the stream

borders. The cattle and bison frequent both the forests and meadows.

Sus scrofa ferus

Cervus elaphus

Megaceros Germanie

Cervus maral (Persia)

Capreolus capreolus

Alces latifrons

Equus mosbachensis

'Trogontherium cuviert

Castor fiber

Sciurus vulgaris

Lepus cuniculus

Arvicola amphibius

= ratticeps

é agrestis

si glareolus

Myoxzus

Mus sylwaticus

Talpa europea

Ursus deningeri

a arctos

a speleus

arvernensis

Felis catus

Lynchus lyne

Lyne cervaria

Canis lupus

“ neschersensis

“< Suesst

Vulpes alopex

Meles taxrus

Mustela vulgaris

- martes

Gulo luscus

Fatorius putorius

Lutra vulgaris

Bison priscus

Bos primigenius

Equus caballus (7 sp.)

FoREST MAMMALS

Wildschwein

Edelhirsch

Riesenhirsch

Reh

Elch

Biber

EHichhornchen

Wildkaninchen

Wasserratte

Nordische Wuhiratte

Erdmaus

Rotelmaus

Deninger Bar

Brauner Bar

Hoblenbar

gem. Wildkatze

gem. Luchs

Silberluchs

Wolf

gem. Fuchs

Dachs

gem. Marden

Edelmarder, Baum-

marder

Iltis

Fischotter

Merapow MAMMALS

Wisent

Urochs

Wild boar

Red deer, stag

Giant deer

Maral deer

Roedeer

Moose (broad-faced )

Horse of Mosbach

Giant beaver

Beaver

Squirrel

Wild rabbit

Dormouse -

Forest mouse

Mole

Brown bear

Cave bear

Val d’Arno bear

Wild cat

Lynx

Eurasiatic lynx

Wolf

Common fox

Badger

Marten

Pine marten ~

Wolverine

Polecat

Otter

Bison

Urus

OSBORN, REVIEW OF THE PLEISTOCENE aA

Fetorius vulgaris . Wiesel Weasel

Sorex vulgaris Shrew

Cricetus vulgaris Hamster

Arvicola terrestris Schermaus

+ arvalis gem. Feldmaus Meadow vole

Of these forest-living animals the giant deer (Megaceros), the cave

bear (Ursus speleus), the Maral deer (Cervus maral), the giant beaver

(Trogontheri1um) and the early Pleistocene species of horse are among

—

PREHISTORIC

AND =

RECENT AN =

FOURTH

GLACIAL

”

PILTDOWN RACE

iti

PITHECANTHROPUS

RACE

eee

ine AN

G a O

y y

a o

Ee Eee ee) ee

ees Se

Warm Temperate Cold Domestic

Fic. 7.—Introduction, succession and extinction of the faune from the five chief

sodgeographic regions

African-Asiatic, Eurasiatic Forest and Meadow, Tundra, Steppe and Desert, Alpine.

the few forms which became extinct during the Glacial and Postglacial

epochs. The great majority of these species survived with successive

subspecific modifications.

It is a very remarkable fact that this true forest fauna of Europe is

frequently found in the same deposits with the “warm fauna” of African-

Asiatic origin. The bison and wild cattle appear in Europe from early

248 ANNALS NEW YORK ACADEMY OF SCIENCES

Pleistocene times, and in late Glacial and Postglacial times they occur as

companions of the mammoth and the woolly rhinoceros.

Alpine Mammals, Cold Fauna.—During the severe conditions of late

Pleistocene times the Alpine mammals were driven down into the plains

or to the lower mountains and hills, and their remains occur principally

during the last Glacial advance. They are represented both in the draw-

ings and in the sculptures of the men of the reindeer or cave period.

Ibex priscus Steinbock Primitive ibex

Rupicapra tragus Gemse Chamois

Ovis argaloides (Altai Mts.) Argalischaf Argali sheep

Arvicola nivalis Schneemaus Alpine vole

Lagopus alpinus Gebirgsschneehuhn Ptarmigan

Steppe Fauna of Russia and Siberia.—Steppe conditions of climate

were rendered possible in Europe by the elevation and extension of land

much farther to the north and northwest than at present. At such

periods all the tempering influences of the Atlantic Ocean were cut off

from northern Europe and helped to give central Europe a cold, dry con-

tinental climate favorable to dust storms.

Boule, Kobelt?® and Scharff?? have agreed in the opinion that the pres-

ence of steppe mammals affords inadequate proof of steppe conditions in

the country. Other authors (Hilzheimer, 1913), however, strongly sus-

tain the steppe-climate theory. The evidence for steppe conditions of

climate has been strengthened in recent years by the discovery of three

successive loess deposits.

The steppe regions of eastern Europe around the Caspian Sea and of

central Asia still maintain this highly characteristic steppe fauna. The

climate is usually one of hot, dry summers with high winds and prolonged

cold winters with sweeping snow storms. The animals are consequently

very hardy. The fauna includes the jerboa, sushk, bobac marmot, dwarf

pica, hamsters, northern voles, corsac fox, the manul, or Pallas’s cat

(Felis manul). Covering the plains are the larger grazing animals such

as the saiga antelopes, wild asses and wild horses (including the Hquus

przewalskwu type). Another animal which probably belonged to the

Steppe fauna is the Hlasmothertum.

Elasmotherium sibiricum* Elasmothere

Equus przewalskii Wild Gobi horse

“ hemionus Dschiggetai Dzeggetai, wild ass, kiang

28 KOBELT, W.: Die Verbreitung der Tierwelt. Gemissigte Zone. Leipzig, 1902.

27 SCHARFF, R. F.: The History of the European Fauna. London, 1899.

°8 GAUDRY, ALBERT, and BOULEF, MARCELLIN: Materiaux pour l'Histoire des Temps

Quaternaires. Trois. Fasc. L’Elasmotherium, 4to. Libr. F. Savy, pp. 83-104, pll. xvi-

xix. Paris, 1876.

OSBORN, REVIEW OF THE PLEISTOCENE 249

Saiga tartarica Saiga antilope Saiga antelope

Alactaga jaculus Gr. Pferdespringer Jerboa

Lagomys pusillus Zwergpfeifhase Dwarf pica, or tailless

hare

Spermophilus rufescens Rotliche Ziesel : Suslik, or pouched

marmot

Cricetus pheus Kl. Steppenhamster Steppe hamster

Arctomys bobac Bobac, or Polish marmot

Myodes lagurus Steppe lemming

Arvicola gregalis Sibirische Zweibelmaus Steppe vole

Canis corsac Corsac wolf

Putorius eversmanni Steppe weasel

Tetrao tetrix Birkhuhn Moorhen

Tundra or Barren Ground Fauna and Flora.2°—Certain members of

the Tundra Fauna adapted to the long cold winters and short summers

of the lands bordering the Arctic Ocean appeared in Europe at the height

ot each of the great glacial advances. The remains of these animals are

always found within or close to the glacial drifts until the Fourth Glacia-

tion when they spread all over France, Germany and Austria. Thus the

musk-ox (Ovibos moschatus) is recorded in the (?) First Glacial ad-

vance of the Forest Bed of England. A tundra fauna including the rein-

deer (Rangifer tarandus) is recorded (Forster, 1913) with the Third

Glacial advance (Upper Mauer sands). An extensive Tundra Fauna also

appears with the Third Glacial, or Riss Stage, in the “Mammutlehm” of

Cannstatt (Koken, Schmidt, 1912, p. 270); this is termed the “Older

Primigenius” Fauna and occurs on the “high terraces” with the older

Diluvium ; it includes the woolly mammoth, the rhinoceros, the horse and

the reindeer. The Tundra Fauna reappears toward the close of the Third

Interglacial Stage (7. e. “Lower Rodent” layer), but the full series of

species characteristic of the Tundra Fauna are not recorded in Europe

until the Postglacial Stage (7. e. “Upper Rodent” layer), when the entire

Tundra list given below is discovered either mingled with the culture

remains of the Neanderthal race of men in Mousterian times or is repre-

sented in the art of the Cré-Magnon men of the reindeer period. The

full or typical Tundra list of the Fourth Glacial Epoch is as follows:

Elephas primigenius Mammut Mammoth

Diceros antiquitatis Wollhaariges Nashorn Woolly rhinoceros

Rangifer tarandus — Ren Barren ground reindeer

Ovibos moschatus Moschusochse Musk-ox

Lepus variabilis Schneehase Arctic hare

Myodes obensis Oblemming Obi lemming, or Siberian

lemming

22 NEHRING, A.: ttber Tundren und Steppen der Jetzt- und Vorzeit, mit besonderer

Berucksichtigung ihrer Fauna, pp. 81-166. Berlin, 1890.

950 ANNALS NEW YORK ACADEMY OF SCIENCES

Myodes torquatus Halsbandlemming Banded lemming

Canis lagopus EKisfuchs = Arctic fox

Gulo borealis Vielfrass Wolverine (glutton)

Fetorius erminea — Gr. Hermelin Ermine

Arvicola nivalis Arctie vole

Lagopus albus Moorschneehuhn Ptarmigan

Asio palustris Sumpfeule

Cygnus MUSICUS Singschwan

Animals like the banded lemming adapted to extreme northerly condi-

tions generally cling to these very obstinately and perish rather than con-

form to an altered environment (Nehring). This species dwells imme-

diately to the north of the region of coniferous forests, among scattered

shrubs of the common juniper (Juniperus communis), the dwarf birch

(Betula nana), the polar willow (Salix polaris) and the mountain dryas

(Dryas octopetala). Thus we may be confident that the lemmings dis-

covered in Pleistocene times in England, France, Belgium and a large

part of Germany are proof of climatic conditions similar to those of the

present circumpolar region. We must conclude that the borders of the

ice caps were surrounded by tundra or barren ground conditions at sey-

eral Pleistocene stages. The lemmings probably dwelt in the immediate

neighborhood of the glaciers. The existing tundras are characterized by

frozen subsoil and the absence of trees or shrubs except along the river

borders.

The reason for associating the woolly mammoth with this fauna is that

the mammoth as depicted by the men of the Postglacial Stage agrees pre-

cisely in its form, its proportions, and its hairy covering with the mam-

moths which have been discovered in the frozen subsoil of northern Si-

beria and are washing out in large numbers along the northern Siberian

and American coast at Eschholtz Bay and elsewhere. At Thiede near

Braunschweig, a classic locality, the lemming remains are associated

with those of the arctic fox, arctic hare, reindeer, musk-ox and mammoth.

Thus the comparison of certain regions of Pleistocene France and Ger- |

many with arctic Eurasia and the barren grounds of northern Asia and

North America is based on the strongest evidence.

MIGRATION THEORY OF FLORAS AND FAUNAS

The principal contributors to the theory of northward and southward

migrations and to the succession of faunas and floras are Nehring (1880-

1896), Woldrich (1882-1896) and Penck (1896-1909). Such alterna-

tion is held by Penck to be demonstrated in Switzerland, where during

the Third or Riss glaciation the woolly mammoth and woolly rhinoceros

ce ee “Anim M -gsebae li fas

) SMR SAUDER A OSE a ce hts a ee

7a = Vpeine't™ wad JIT?! * 9 ieee (sects re : ‘fnatoqaln +

udieAUAT WO alt, ee! rust eek, pease are aes eTothqeimsi

GAG.

eS ee ae mes ‘,

-- : crs Be Sn Sea

+ wee = TS RE I

2 a

ven rinw the F :

ven : a a Abbe ayadanys anid ' :

Osa aIsaayiasT* ia , ig de Ries a be

aie StTTITA .ARGUT LENGE OF 2 A REEDS SY | Dasvbissad

“As ar? disperses} OOU TEE.

ate AA. SHATTER: > OYE e oW. poy: 4 es

‘woe Kcitt awh Trt i. Weutieaaed 7s "BON DTD “20% Hongo mibdf 30

ring t ne Dein — Amdvalot & BOTAN Bend

one successive series IN’ BWOBING sousd

w *

su con = ae

= m * of different faun® EwsnTybonrig i ara ae

rmeratire and of flodeimern of Wee _ ebfslon sit 498k > Shoilst- base

Shey ese aVeagree 2 ee bpetebipes pee as anwpil 5aEA *

iiouley when Cee er ad Bist as Saat 8 papt

a of i fe “A RIPG aie pers bite sie Seite : Par 3

", z a , a ‘faucs ame. ate 1 =

li yy erguact ys Eee Ss yi “Ty eS IT

pent Re : ERS eye Bee ate © _susiToge anerd ae =

Same generic MD ERE Seek Ss oe eet aio Me Sitn 28

TL the Case OF Lne-4 oe 3 ; meek __stingts awrinioe% | re ; ees |

GR) aaioumM I |. easgist ainnd FIRE

nh eH a pete Se aint ST, SEs pore ai

ely, that = 5 aROd eatolh

at St ee EE i PES

“enor ited -e9oLk

: * | ees Tustira (2a EE a eo ee

Siileenive) Senge tu 98H ote hee eee

aby. Mauer, FONG MOAI OQOWT ze |

; nn ; _Heroybsts patnd

emer — aiog OLED

ciaaeee ae. *eiehonsiiaoue espe

es POS =. a ees $ wed ae

a, eee

ino Fs =:

tila t

nely

fs ae B

RECENT : L Ili Pure Forest

PREHISTORIC of ] Mrapow Fauna

POSTGLACIAL

i Il “REINDEER PeERIOD,”’

(Severe climate) REINDEER Perron,

TUNDRA, STEPPE,

Vy Asawa — “a ALPINE AND MEADOW

Fau FAUNA

(Steppe climate) :

3rd INTERGLACIAL

III GLACIAL el

I MINGLED AFRICAN-

2nd INTERGLACIAL

De ASIATIC AND

ForEsTt-MEADOW FAUNAS

II GLACIAL

1st INTERGLACIAL

I GLACIAL

GEOLOGIC AND CLIMATIC FAUNAS

STAGES

HABITATS

Diagram illustrating theils covered all except

the areas close to the ice. Thteppe forms covered

Europe. .

31 Hyena speiea and Felis leo spel@a, 1

Present

Cold Climate

Temperate climate excepting near the glaciers

OSBORN, REVIEW OF THE PLEISTOCENE 951

first appear in Europe, only to be succeeded in the Third Interglacial or

Riss-Wiirm Stage by the reappearance of the warm African-Asiatic fauna,

including the straight-tusked elephant (£. antiqguus) and the broad-

nosed rhinoceros (D. merckw). Theoretically this warm fauna was again

driven out during the Fourth or Wurm glaciation by the reappearance of

the woolly clephant and the woolly rhinoceros. Penck*® observes, as

noted above, that we cannot hope to trace a continuous evolution of forms

during the Pleistocene because we are not dealing with the development

of one successive series in one locality but with the cycical evolution of a

number of different faunas compelled to migrate because of alternations

of temperature and of flora, the mammals disappearing and returning at

intervals too brief to allow of any marked evolutionary changes. Herein

hes our difficulty when we attempt to distinguish between the Tundra

Fauna of the Third and Fourth glaciations and the Forest Fauna of the

corresponding interglacial stages, because the faunas return not only with

the same generic but with the same specific types, as is especially illus-

trated im the case of the mammoth (LZ. primigenius) and the giant deer

(Megaceros).

_ Resident Theory.—Another theory is presented in the accompanying

table, namely, that during the First, Second and Third Glacial Stages

the climatic conditions of Europe were temperate except in the immediate

regions surrounding the glaciers. It was these glacial border regions

which attracted the tundra fauna of the north, the reindeer and the

woolly mammoth. Supporting this theory is the fact that the tundra and

steppe fossils are found only in proximity to the ice caps and glaciated

regions. It is not until the Fourth Glacial Stage and the Postglacial that

the general climate of HKurope was so severe as to cause the gradual emi-

eration and extinction of the African-Asiatic fauna. At this stage re-

mains of the tundra and steppe mammals occur in all parts of Europe,

including the Pyrenees and Cantabrian Alps, but only partly extending

into Spain.

Latitude and Altitude——In considering the distribution and migration

of the mammals throughout the Glacial Epoch, we must constantly keep

in mind the differences of latitude and of climate which prevailed then

as now between Italy, Spain, southern and northern France, Germany

and Belgium: also the differences of altitude as between the lowlands of

the rivers Rhone and Dordogne and the highlands of the Alps and other

mountains. Italy had a more moderate climate than central Europe: the

reindeer seems never to have found its way there, yet a lowering of tem-

80 PencK, A.: “Die alpinen Eiszeitbildungen und der prihistorische Mensch.” Arch.

Anthropol., N. S., Vol. I, No. 8; pp. 78-90. 1°04.

est Meadow Mammals

Tasty IV.—Distribution of For

88S

a. Oe

Return

2 dispersal

of } forest

of Alpine Mauna to the Mountains. Wid¢ Temisphere

and Meadow Fauna oyer entire Northern F

TIL Purr Worrsp

MEADOW Fauna

RECENT

PREHISTORIC

Present

Cervus elaphus

POSTGLACIAL

II “REINDEER Prrrop?’| 2

Retr 1 i hyrardsand Se CUTE OLS TUNDRA, STEPPE oe

(Severe climate) etreat of Tundra and Steppe Fauna. Wide sout | in Burope NDRA, STEPPE, E

wesstward dispersal of Tundra, Steppe and Alpine Faunz of Tundra Megaceros ewryceros ALPINE AND Mpapow =

3 e to the Pyrenees, Alps, northern Spain and Italy. Arrival Ursus arctos Fauna

IV GLact Favina. Arriyal of Asiatic Steppe Fauna.

Bison priscus

Bos primigenius

aie ae -hward retrea Alces machlis

Return of Warm and Tem- Hxtinction or sout parame f va

8rd INTERGLACIAL perate Wauna E of African-Asiati: ~ BHawus caballus typicus

Cold

(Steppe climate)

lea Felis catus

eee ee | «ne

Vulpes alopea

Hyena crocuta spe

8rd Tundra (Rangifer = B.

Aas f Felis leo spelea™

prunigenius) fauna in north-

III Guacran ern Germany and France, Dicerorhinus mereh ™ Ursus speleus

Ge of Alps and Pyrenees, Blephas antiquus Felis catus

Blephas trogonther “ Sciurus vulgaris

Hippopotamus maj 2” Canis lupus I Minetrp Aprrican-

Return of Warm and Tem- Cani = Sus scrofa

2nd INTPRGLACIAL perate Mauna ALIS ATS ASIATIC AND

Macacus Meles tarus

Felis pardus Alces latifrons Forest-Mrapow Faunas

2nd Tundra (Rangifer — HB. ee

Castor fiber

Trogontherium cuvieri

Lutra vulgaris

Gulo luscus

Hquus mosbachensis

mimigenius) fauna in N. Ger-

I Gracia I Sh iat

I tiany, Mauer, Steinheim, etc.

Dicerorhinus etrusc Us

Macherodus latide

Bquus stenonis

Return of Warm and Tem- Hyena striata

ist [NDTERGLACTIAL jerate Fauna

Temperate Climate excepting near the glaciers

Blephas meridional 8 Ursus deningeri

Feet Ursus arvernensis

Survival of Plioce aa ane Tene Cervus sedgwicki

— ist Tundra (Ovibos) fauns, dnvasions oF jA*rican “Ss Cervus carnutorum

I Gracrar 1 Great aritain Fauna

[Prccrca SS a

aE fe : fRICAN- DURASIATIC Formst

in igrations in- “Warm” Al ; =;

vly TUNDRA migra 4 PAUNA

GEOLoGIO aND Crimatic Siem Europe and cold ASIATIC FAUNA, aes Reena a x TrequeAtn BAUNAS

Sracrs jons of Britain, Alps and or migrating into eed ts

T S d Sheltered Re- Cool temperate meadows and for-

‘enees Temperate an

gions, remote from the Ice Caps ests. Regions bordering and re- HAsirars

\— ae! 2 mote from the Ice Caps

(RCUMGLACIAL HABITATS AND ee

GLACIAL MicRANTS Trmprrate NON-GLACIATED HABITATS AND INTERGLACTAL

Micrants

Dia illust th thor’s theory that until the Fourth Glaciation the African-Asiatic-Murasiatic Forest Meadow Mammals coyered all except

gram illustrating the author

the a i] to th The tundra forms were confined to the ice borders uitil the Fourth Glaciation, when the tundra and steppe forms coyered

reas close to the ice, e tundra

Durope.

in the Colc Reindeer period of IV Glacial and Postglacial times,

" Hyena spelea and Felis leo spelwa, the sole survivors of the African Asiatic eure

592 ANNALS NEW YORK ACADEMY OF SCIENCES

perature in Italy is mdicated by the fact that the Alpine mammals such

as the marmot (Arctomys marmotta), chamois (Rupicapra) and steinbok

(Ibex) came down to the plains.** The “old elephant” (£. antiquus)

fauna appeared in Italy earlier than in northern Europe. Similarly the

hippopotamus survived in Italy longer than in France and Germany, so

it is not surprising to find its remains associated with those of the broad-

nosed rhinoceros (D. merckw) in a cave near Mentone on the French

Riviera which belongs just prior to the Postglacial period of Aurignacian

culture corresponding with the period immediately succeeding the Fourth

glaciation.*® Italy was a forested country at the time that central Europe

was tundra-like or steppe-hke.

Culture Stages—Under all these varying conditions of climate human

evolution progressively advances, and where the “species” of lower mam-

mals fail us the successive human culture stages enable us to sharply

distinguish the intervals of time.

Repetition of Loess Depositions.—According to Wiist** (p. 229) the

First (1) and Second (II) Glacial and the First Interglacial Stages (1st)

are represented in the single northern (Saxonian -++ Scanian) glaciation

of Thuringia. The successive depositions of loess therefore occurred as

follows:

Postglacial Stage, the youngest loess. Postglacial terraces.

IV (WURM) GLACIAL STAGE, equivalent to northern glaciation III.

3rd (Riss-Wiirm) Interglacial Stage, younger loess. The lower terraces

of the 2nd (i. e., 3rd Interglacial).

III (RISS) GLACIAL STAGE, equivalent to glaciation II of the north.

2nd (Mindel-Riss) Interglacial Stage. older loess. The higher and middle

terraces of the First (7. e., 2nd) Interglacial.

I-II (GUNZ-MINDEL) GLACIATION, equivalent to glaciation I of Thuringia.

First GLActIaL STaGE—GUNz, Scanran, NEBRASKAN

The First Glacial Stage is believed to have been nearly contempo-

raneous in Europe and North America. It is estimated (Penck) that

the various ice caps reached the climax of their advance 520,000 years

ago and that the snow line in the Alpine Region descended 1,200m. lower

than the present snow line. In the north of Germany and in Sweden

2IsseL. A.: “Liguria geologie e preistorica.’”’ Ref. by Boule in L’Anthrol., pp. 602-

604. 1893.

33 PENCK, A.: “Die alpinen LEiszeitbildungen der prahistorische Mensch.’ Arch.

Anthropol., N. S.. Vol. I, No. 8, pp. 78-90. 1904.

% WtstT, EwaAbp: “Die plistozanen Ablagerungen des Travertingebietes der Gegend

yon Weimar und ihre Fossilienbestiinde in ihrer Bedeutung fiir die Beurteilung der Kli-

maschwankungen des Biszeitalters.” Zeitschr. f. Naturwissensch.. Band 82, Heft 3-5,

pp. 161-252. Leipzig, 1910. Published Mar., 1911.

OSBORN, REVIEW OF THE PLEISTOCENE 953

the deposits of this glacial advance are known as the old “Diluvium” or

the “Oldest Drift;” the advance is termed the Scanian by Geikie. In

the Alpine Region it has been termed the Ginz by Penck and Briickner,

and the drift deposits have a general thickness of 30m. At about the

same time a great ice cap was formed in British North America west of

Hudson Bay from a centre known as Keewatin which sent its ice sheets

into Iowa and Nebraska. The resulting Nebraskan deposits, consisting

largely of compact boulder clays, are often thickly set with woody ma-

winupeg

puital0

> of Li

Few?

30 West Longitude 80 from Greenwich

Fic. 8.—Chief centers of North American glaciation

Keewatin, Labradorean, Cordilleran. After Leverett.

terial gathered from forests of spruce and other coniferous species that

indicate the development of a cool temperate flora in advance of the

glaciation.**

It does not appear that a glacial cap of any considerable extent was

formed in Great Britain; but Geikie®* shows that along the British coast

% The above correlation is presented chiefly on the authority of Penck and Leverett

(op. cit., 1910).

36 GWIKIE, JAMES: The Great Ice Age. 2nd Ed. London, 1877.

254 ANNALS NEW YORK ACADEMY OF SCIENCES

in Upper Pliocene times cold conditions began to manifest themselves

in the Pliocene Red Crag and continued with increasing intensity during

the deposition of the post-Pliocene Chillesford and Weybourn Crags,

which mark a culminating time in which the sea abounded in Arctic

moiluses, and this may represent the first glacial stage in Britain.

Fic. 9.—Principal mammal deposits and culture stations of the Pleistocene of Europe

1. Forest Bed of Cromer (Norfolk). Sables de 2. St. Prest near Chartres (Eure-et-

Loire). 3. Malbattu (Puy-de-Déme). 4. Peyrolles (Bouches-du-Rhéne). 5. Solhilae

near Puy. Clay deposits of 6. Durfort (Gard). 7. Cajare (Lot-et-Garonne). 8. Val

d’Arno (Tuscany). 9. Leffe near Bergamo (Lombardy). 10. Rixdorf near Potsdam

(Brandenburg). Gravels of 11. Siissenborn near Weimar. Sands of 12. Mosbach in

northern Baden. Fresh-water deposits of 13. Clacton (Essex). Sands of Mauer near

14. Heidelberg (western Germany). 15. Chelles on the Marne, near Paris. 16. St.

Acheul (Somme). 17. Ilford and Grays Thurrock (Essex). Lignites of 18. Diirnten

and of Utznach, near Ziirich. 19. Taubach near Weimar. 20. Wildkirchli cave on Mont

Santis (eastern Switzerland). Tuffs of 21. the Tiber Valley, near Rome. Caves of 22.

Neandertal, near Diisseldorf (western Germany), 23. Spy, near Amur (Belgium), 23a.

Krapina (Croatia), 24. Chapelle-aux-Saints (Corréze). Caves and alluvial deposits of

25. Ternifine (or Palikao) near Oran (Algeria), 26. Pointe Pescade, near Algiers (Al-

geria). 27. Prince’s Cave (Monaco). Sandy clays of 28. Véklinshofen (Alsace). 29.

Saalfeld (Saxe-Meiningen). Travertines, etc., of 30. Gera, Jena (Saxe-Weimar). 31.

Leipzig (Saxony). 32. Solutré, north of Lyons. Loess of 33. Wiirzburg (Bavaria). 34.

Thiede near Braunschweig (Prussia). Cave of 35. Montmaurin (Haute-Garonne). 36.

Chateauneuf-sur-Charente (Charente). Caves of 37. Schweizersbild near Schaffhausen,

and Kesslerloch near Thayngen (northern Switzerland). Remains of lake dwellings at

38. Wauwyl (Lucerne), 39. Robenhausen, south of Lake Pfiffikon, 40. Concise on Lake

Neuchatel (Switzerland). Peatbogs of 41. Hassleben. near Weimar. Travertines of 42.

Langensalza (Erfurt) in central Germany. Caves of the 43. Island of Malta, 44. Island

of Crete, 45. Island of Cyprus.

OSBORN, REVIEW OF THE PLEISTOCENE O25

fey |

First INTERGLACIAL STAGE

-NORFOLKIAN, AFTONIAN, GUNzZ-MINDEL

Immediately after the deposition of the Weybourn Crag in Essex there

is evidence of a climatic reaction because the overlying deposits, both

estuarine and fluviatile, of the so-called Forest Bed of Cromer (Norfolk)

contain a flora and mammalian fauna of warm temperate type which

contrast strongly with the assemblage of the northern and Arctic mol-

luscs in the subjacent deposits of the Red, Chillesford and Weybourn

Crags. From this classic locality of Norfolk the First Interglacial Stage

has derived its designation Norfolkian. It is also known as the Cro-

merian.

The first cold period or glacial advance was succeeded both in Europe

and America by climatic conditions milder than the present. In the

Alpine region Penck and Brickner have found evyidences.of a long

Giinz-Mindel interval of time, estimated relatively at 100,000 years,

which separates the chmax of the first or Giinz glaciation from the sec-

ond or Mindel, but they believe that this First Interglacial Stage was

much shorter than the Second Interglacial. Owing to the warmth of

the climate the snow line of the Alps is believed to have risen 300m.

higher than at the present time. The deposits of this stage include the

‘“Paludinenbank of northern Germany; these fresh-water deposits near

Berlin are of true interglacial time, since they he between the deposits of

two Glacial advances. The most abundant species, Paludina alluviana,

has its present habitat far to the south on the borders of the Black Sea.

In general the other fresh-water and land molluscs belong to modern

species common to the same region to-day, indicating that climatic condi-

tions were not greatly dissimilar from the present. The plant remains

include the mosses and conifers, also indicating a climate similar to the

present, but they probably do not afford evidence of the higher ranges

of temperature which may have occurred at other times and places during

this First Interglacial Period. In America the deposits of this First

Interglacial Stage, known as the Aftonian, are widely distributed and

yield a rich mammalian as well as a land molluscan fauna and abundant

plant remains which generally indicate a temperate to warm climate.

HUMID FOREST CONDITIONS

A striking characteristic of this interglacial stage is the evidence of

prolonged humid conditions of climate and abundant forestation favor-

able to forest-living mammals. There is no evidence either in the north

of Germany or in the northern United States of loess deposits such as

occur during the Second and Third Interglacial Stages and are believed

to be due to recurrent arid conditions and dust storms.

256 ANNALS NEW YORK ACADEMY OF SCIENCES

FLORA OF THE FIRST INTERGLACIAL PERIOD

In the “Forest Bed of Cromer” on the eastern coast of England the

arrival of Abies is significant because although known in’ Miocene times

in the Arctic region of Grinnell Land this is the first appearance of the

fir tree in central Europe; the fir is also found in the interglacial lignites

of Switzerland and has since constituted an important member of the

European forests. Including the fir, all the trees composing the forests

in the region of Norfolk belong to living species, such as the maple, elm,

birch, willow, alder, oak, beech, pine, spruce, still indigenous to this re-

gion, latitude 52° 40’ N. A notable fact in examining this flora of the

Norfolk and Suffolk coasts of England is its correspondence with the

modern flora in spite of the immense period of time that has elapsed and

the great changes in climate during which all these plants were driven

to the south and again permitted to return. “However,” continues

Reid,*" “though very similar, we find in the fossil flora several exotic

species which give it a slightly different character and we notice also the

absence of several modern forms.” From this tree flora Reid concludes

that the climate of southeastern England was nearly the same as at pres-

ent, but slightly warmer.

The flora of Durfort (Gard) in southern France is associated with re-

mains of the southern elephant (. meridionalis), the etruscan rhinoce-

ros (D. etruscus) and the Pliocene horse (Hquus stenonts).*® It in-

cludes numerous plants of species now represented in the Caucasus, Per-

sia, southern Italy, Portugal and Japan. Again, in the interglacial for-

ests of Moret (Seine valley) we find the fig (Ficus) and the Judas tree

(Cercis), indicating a mild temperature. The tree flora of France like

that of Norfolk thus indicates somewhat warmer conditions of tempera-

ture than prevail at the present time, a temperature of 4° of latitude to

the south.

MAMMALS OF THE FIRST INTERGLACIAL STAGE

For our knowledge of the mammalian life of the Forest Bed of Nor-

folk and contemporaneous deposits of France we are indebted principally

to Dawkins (1880, 1883), Newton (1880), Gaudry (1893), Boule

(1902), and Pohlig (1907). Dawkins many years ago (1883, p. 579)

estimated the ratio of living, extinct and newly arriving mammals in the

Forest Bed as follows:

37 REID, C., and Reip, E. M.: “The Pre-Glacial Flora of Britain,” Jour. Linn. Soc.,

Botany, Vol. xxxviii, pp. 206-227. Jan., 1°08.

3% GauDRY, A.: L’Eléphant de Durfort. Paris, 1893.

OSBORN, REVIEW OF THE PLEISTOCENE 95!

Survivalisperom phe: PHOCENE J 3 depots 12s ee ~ ss ee 11 species

NewCOMmers, TOrmMs NOW Extinet-< ..)...722% 5-5 0,2 ae 6 -

ING W COMERS) LoOrms StH Divine . Sac sce ate ela wien e eewiens AN es

The specific determinations of many of these animals are based upon

very incomplete materials and await careful revision; and upon this

closer study will depend also the correlations between the various First

Interglacial faunze of Great Britain and the continent. From our pres-

ent knowledge the following faunistic correlation may be made:

Northern Life. Forest Bed of Cromer, Norfolk, England (Fig. 9, 1), typical

of the northern life.

Sables de St. Prest (Eure-et-Loire), France, (2) typical of the

central life of France.

Malbattu (Puy-de-Dome), France, (3) typical of the central

life of France.

Southern Life. Peyrolles (Bouches-du-Rhone), France, (4) typical of the

southern life of France.

Solilhac, near Puy, southern France, (5) typical of the south-

ern life.

Durfort (Gard), southern France, (6) typical of the southern

life.

Cajarc, Lot-et-Garonne, (7) typical of the southern life.

Val d’Arno (upper, or Pleistocene deposits), northern Italy,

(8) typical of the southern life of Europe.

Extinctions—The survival of a number of Plocene mammals into

this period has led certain paleontologists, such as Boule, to place the

First Interglacial fauna and the Glacial stage which preceded it in the

Pliocene rather than in the Pleistocene Epoch. The true Upper Plio-

cene fauna, however, is characterized by a number of distinctively Plio-

cene mammals, especially the primates, mastodons, antelopes, gazelles

and tapirs. None of these animals have been found in the Pleistocene of

Europe; all had become extinct.

Survwals.—This First Interglacial fauna does include, however, a

number of -survivals from the Pliocene, such as the sabre-tooth cats

(Macherodus), the polycladine deer (C. sedgwicki), the more primitive

dicerorhine rhinoceros with brachyodont teeth (D. etruscus), the primi-

tive Upper Pliocene horse (Hquus stenonts), the southern elephants (F.

meridionalis), the hippopotami, the roedeer (Capreolus capreolus), the

giant beaver (Trogontherium), the Auvergne bear (Ursus arvernensis),

the wild cattle (Leptobos).

It is noteworthy that Pohlig considers the 2. meridionalis of this stage

to be of more recent type than the true Pliocene type of Italy described

by Nesti; hence he terms it #. (mertdionalis) trogontherii. It is also

258 ANNALS NEW YORK ACADEMY OF SCIENCES

noteworthy that the polycladine deer (Cervus sedgwicki) do not reappear

in any of the subsequent Pleistocene formations of Hurope.

Arrivals—Among the new arrivals in the Forest Bed of Norfolk are

the earliest members of the giant deer race (Megaceros) which continues

into Middle Pleistocene times in Europe. We also note in the Forest

Bed the presence of a form (Caprovis) intermediate between the goat

and the sheep, as the name indicates, and most closely resembling the

moufflon of Sardinia. Among the rodents the large beaver Trogonthe-

Hig. 10.—Giant deer, Megaceros, of the British Pleistocene

From a skeleton found in the Irish peat bogs. After original by Charles R. Knight in

the American Museum of Natural History.

rium cuvrert succeeds the smaller ancestral species (7. minus) first ob-

served in the Pliocene of the Red Crag. The giant hippopotamus (4.

major) is certainly recorded in this region of Great Britain as well as to

the south in Italy.

Among the proofs of a northerly climate is the first occurrence of the

musk-ox (Ovibus), which is attributed by Dawkins* to the Forest Bed

deposits.

°° DAWKINS, W. Boyp: ‘On the Alleged Hxistence of Ovibos moschatus in the Forest-

bed, and on its Range in Space and Time.” Quart. Jour. Geol. Soc. London, Vol. 39,

pp. 576-579. 1883.

OSBORN, REVIEW OF THE PLEISTOCENE 259

Among the animals attributed by Dawkins to the Forest Bed fauna

which Newton*® considers of doubtful reference are the hyena, the true

cattle (Bos primigenius), the red deer (Cervus elaphus), the moose

(Alces latifrons) and the giant deer (Megaceros). ‘These animals are,

however, certainly recorded in France (Cajarc) with the exception of the

moose (Alces).

The presence of deer (Cervide) in great numbers and representing

many different phyla is one of the most distinctive features of First In-

terglacial times. There existed numerous and varied forms of deer life

both in Great Britain and southern and western Europe, attesting the

presence of forests. They belong to several, probably to as many as five

distinct phyla. Among these the polycladine, or “many-branched” deer

so distinctive of the Upper Pliocene of the Val d’Arno now make their

last appearance in Europe as Sedgwick’s deer (C. sedgwickv) of the For-

est Bed, with remarkably complex antlers closer to the C. dicranius of

the Val d’Arno. A second Phocene European phylum is that of the roe-

deer (C. capreolus). A third phylum, numerous and highly diversified,

is that of the giant deer (Megaceros) which is represented by a variety

of species (M. verticornis, M. fitchu, M. dawkinsi). There is some doubt

whether the true “deer of the Carnutes” (C. carnutorum) occurs here.

As above noted Newton is uncertain whether the stag (C. elaphus) has

been truly recorded in the Forest Bed. Neither the true fallow deer

(Cervus dama) nor the reindeer (Rangifer tarandus) has been recorded

there.

This Forest Bed fauna as a whole is an outlier of the Asiatic-African

group with a strong northerly Eurasiatic forest element intermingled.

We observe that browsing, forest-living and fluviatile types predominate.

Among the forest-frequenting carnivores are the wolverine (Gulo), the

otter (Lutra), two kinds of bear (Ursus), the wolf (Canis), the fox

(Vulpes), the marten (Mustela) and a true feline (Felis) in addition

to numerous representatives of the sabre-tooth tigers (Macharodus).

All the above are true Eurasiatic forest types from north-temperate lati-

tudes. Among the forest-living browsers also is a large boar related to

Sus scrofa, the primitive browsing rhinoceros (Dicerorhinus etruscus)

with short-crowned teeth, while in the rivers disported the giant hippo-

potamus (H. major). Among the grazing and meadow-living forms

Bos or Leptobos is represented in the Forest Bed. There are also two

species of horses including a lighter form resembling the FH. stenonis

40 Newton, ©. T.: “Notes on the Vertebrata of the Pre-Glacial Forest Bed Series of

the East of England.” Geol. Mag., Vol. vii, Pt. I, Carnivora, pp. 152-155, Pt. II, Car-

nivora, pp. 424-427, Pt. III, Ungulata, pp. 447-452, Pl. xv. 1880.

960 ANNALS NEW YORK ACADEMY OF SCIENCES

cocchi of the Val d’Arno, and a heavier type of horse, probably forest-

living.

Cool estuarine conditions are represented by the presence of the walrus

(Trichecodon), the seal (Phoca) and the cormorant (Phalacrocoraz).

The chief members of the mammalian assemblage of the First Inter-

glacial Stage in Britain and France are as follows:

Southern elephants

E. meridionalis (trogontherii)

Dicerorhine rhinoceroses

D, etruscus

Primitive horses

E. stenonis, EF. caballus fossilis ?

Hippopotami

H. major

Polycladine deer

C. sedgwicki

Roe deer

C. capreolus

Giant deer, Megaceros

M. dawkinsi, M. verticornis

Primitive cattle

Bos primigenius

Giant beaver

Trogontherium cuvieri

Sabre tooth cats

Macherodus ? cultridens

Wolverines or gluttons (in Great

Britain)

Gulo luscus

Bear of the Ursus arctos type

Otters and martens

Wolves and foxes

Walrus (in Great Britain)

Trichecodon huxleyi

(In Italy only)

Straight-tusked mammoths

EH. antiquus

(In France)

Bison

Bison ? priscus

Stag

Cervus elaphus

mus (H. major), the giant beaver (Trogontherwm).

the first remains attributed to the bison (B. priscus).

FIRST INTERGLACIAL OF FRANCE

The Lower Pleistocene deposits of

France, which are approximately of

the same age as the Forest Bed (Nor-

folk), are those of St. Prest (near

Chartres), of Durfort (Gard), of

Sollhac (near Puy) and the re-

cently discovered phosphorite de-

posits of Cajare (Lot-et-Garonne)

(Fig. 9,7). All these beds are con-

sidered by Boule as of Upper Plio-

cene age; they are placed in the

transition period between Pliocene

and Pleistocene times by Harlé and

Stehlin** as well as by Depéret (St.

Prestien ). }

We are disposed to consider all

these deposits of approximately the

same age as the Forest Bed, namely,

Early Pleistocene. In none of them

have remains of primates, masto-

dons, tapirs’ or other characteristic

Upper Pliocene mammals been

found. On the other hand, these

beds contain several forms surviving

from the Plocene, such as the south-

ern mammoth (fH. mertdionalis),

the etruscan rhinoceros (D. etrus-

cus), a primitive species of horse

(EH. stenonis), the giant hippopota-

In France occur

In Durfort was

41 HARLE, E., and STRHLIN, H. G.: “Une Nouvelle Faune de Mammiféres des Phos-

phorites du Quercy.” [Cajarc.] Bull. Soc. Géol. France, Ser. 4, Vol. IX, pp. 39-52. 1909.°

OSBORN, REVIEW OF THE PLEISTOCENE 961

obtained the skeleton of the southern mammoth described by Gaudry.*

The giant deer (Megaceros) are represented at St. Prest by the species

CO. carnutorum, or “deer of the Carnutes.”’

The most recently discovered fauna of this age is that of Cajare in the

northern part of the Rhone basin. Harlé and Stehlin consider the Cajare

fauna as transitional between that of St. Prest and Durfort and that of

Montmaurin which we attribute to Second Interglacial times. The mam-

moth (2H. meridionalis) is of a more recent type than the Upper Pliocene

form of Italy. The bison is a very

large animal. Especially important

is the fact that these authors posi-

tively identify here remains of deer

which are related to the stag or red

deer of Europe (Cervus elaphus),

leaving the species doubtful. There

is also at Cajarc a large badger

(Meles taxus) which is character-

istic of early Pleistocene times, and

a small wolf (Canis) no larger

than the jackal. There is also a ?

small species of sabre-tooth tiger

(Macherodus).

Southern Elephant (Hlephas

meridionalis ).—The “southern ele-

phant” is Faulkner’s “pre-Glacial

variety of the mammoth.” It be- '

longs to the same general group as

the other mammoths (LH. trogon-

theru, EH. primigenius) and the iia tD

Columbian mammoth (£. colum- So ae as.

bt) of America. The southern ele- erg tie id dS

phant of First Interglacial times

belongs to a somewhat more advanced type than that of the Upper Plio-

cene of the Val d’Arno; the Forest Bed specimens are, in fact, attributed

by Pohlig*® to his species H. trogontherii. This mammoth series is dis-

tinguished both from the African (Lozxodonta) and the Indian (Fuele-

phas) elephants by the peculiarly flattened and concave forehead and the

high, peaked cranium. It is probably descended from £. planifrons of

the Pliocene and in tooth and skull structure it resembles the HL. hysu-

Fig. 11.—WMolars of Pleistocene elephants

42GaupryY, A.: L’Bléphant de Durfort. Faris, 1883.

8 PoHLIG, H.: “ther Hlephas trogontherii in England.’’ Monatsber. Deutsch. Geol.

Ges., Vol. 61, No. 5, pp. 242-249. 1909.

7 - f iL. = : ee 3 Sey S| Be qe See bok L = a fags =

4} -- = Aes — = = eS wee Pe

e 2 Qe ; ~ | ;

Ss UF LHC bs PROCES OLWalias STR Ia. HuUS LW tLISt CL Stlastcs

# 44 wee mama ar — es eo oe Wereswmwse S249 Sean at

ea < = 2 k ; G , e

e235 Bii- arn RECESS E SE G Ss Et ZAESCP WS Fe. S2SQ, SSECFE DL RPS SSSECE ET Ls PLA_ULE § §55™- Gb

= 7 —_. = | eters am +h Teams = =o j 2 —. ne

ge cf Qe = e ee e ‘ 4

EER LEO, VE Le CEES FY SEF LESS SEPA EES U Belize at 3 ZSSQF, CLESLELE LESSIG

> tery ions TES eS ooImes teeth wt thick Pam. + athear nore nr;

Fu! SPW. VIELE S FE EEECEEEE™” ERLE WELLER LELEIEN CERGISIT ET, LET PLEEQU ER SESRPER GF a

: = = a - - Aree ~ =

Ra sy cia cre Re neo ot Othe BB aOrect a" a+ Dorfort and tm +ne PI BIS

SF ECRELVO Bta@a OC. VUULULT ESE S85 LEI § CFEC SL phHec. at CRE SSPE RL. GEETE EEE LEI ICIS

= : ~ =p -— =- : == = ~- = = = =

ee | ae EE a Ay a ] Wes and Aisthmomiched hy danta mlatec o-

LOcene GEPOSILS OL LHE al @ ATMO, and distumevsuisned DV Gental plates OL

bs 3 - £

) ea ee wi Pes Set = 4 +}. at pe = aH

e 2 e ra 3 Qe = > e ge o 2 e @e

LELREEERE SE Cie bie’ The FALLOT §S Salli LY VE LUG iSitia’® LV Ui “NCSL We fie?

t Se - 2 —— = = ee SL SSeS be So =St = joes eS ee Se

— —_— a _ ~ _ i ~- = :

SQUPULESUEC NE LEI SPUUICS cei LEEE ESIGLLCE EZ SPOR EEER LV EM ECAPULEEEGCDS EEVOSLE aePit,

Fié > __ Skelei 7; Ele I= me onatis 7 Duri t

Gallery of Palzontology in the Museum d Histoire Naturelle. Jardim des Plantes, Pans

After Gaudry

The southern elephant is best known from the Nagnincent specimen

phat a | ee Ee i 15 oe Ss Sees oe ead =e = 413) ii es “< tad al

aris Museum. he height at the shoulders is 3.83 m.. or 12 f ft., 9-1/5

m. This animal was much taller than the true mammoth (£. primt

enius) which tirst appears chiery in the I hird Glacial Stage. he tusks

were shorter and less bent. We may mifer from its original warm-tem-

perate habitat that it was partially hairy but not covered with wool like

his of the great

as the extinct forms are

be]

f=)

a |

faa

bef

—_

faa)

ine

e\)

frond

hel

is")

lar}

emt

—)

fa a)

iva]

jade

iv 2]

fonda!

honby

i=]

Lae |

OSBORN, REVIEW OF THE PLEISTOCEND 263

estimate the height of these animals from the skeleton. In almost all

cases of mounting these animals in museums the tips of the dorsal spines

are unduly elevated above the superior spine of the scapula, which gives

an exaggerated estimate of the height.

General estimated height at

shoulder

English measure Meters

L. imperator, the imperial mammoth of North America,

Mat CISUOCONG Yat ne we he Mo atte. ssa. veloe be viele aes 13’ 6”-14’ 4.2672

The straight-tusked elephant. #. antiquus of Europe.

fe recuimnated. (POnie, Piulorim) ate... o. ees b's ws ee 1G 4.8768

The southern elephant of Europe. £. meridionalis....... BO Bie! ies 3.8862

The largest living African elephant, L. africanus....... 11’ 3”-11’ 4” 3.4544

The Columbian mammoth of North America, EF. columbi. 9’-11' 3.0028

tment elephant of Asia, H. Mdicus..........00550 9’ 10”-10’ 3.048

The true, or northern mammoth, EL. primigenius (teste

1 ELS) See “et. oc REL Re ee eee 9’ 6”-10' 6” 3.2004

The living pigmy elephant of the Congo, L. cyclotis

Pe EC SEE TOPNAGAY ). ois ek ccc cea ce cee eee ee 4’ 5” 1.3462.

The dwarfed elephant of Crete (EF. antiquus creticus),

Malta (#. antiquus melitensis) and Cyprus.......... ay 1.524

The American mastodon, Mastodon americanus......... G64 2.8956

Hippopotams.—The hippopotamus is the invariable companion of the

southern elephant and later of the Hlephas trogontheri; it survived even

in northern Europe until the middle of the Third Interglacial Stage.

Outside of Asia the oldest hippopotamus remains known are from the

Lower Pliocene of Gravitelli in Sicily;** it is compared by Seguenza

(1902) with the H. swvalensis of India but is clearly distinct from this

species. The H. hipponensis of Gaudry is confined to the Middle Plio-

cene of North Africa and throws no light on the phylogeny of the hippo-

potami. These animals appear in the Lower Pliocene of India and of

Sicily and in the closing Phocene of Italy, India, and North Africa and

perhaps China; in the Quaternary they spread into Java, India, Mada-

gascar, Africa from north to south, the Mediterranean Islands, and from

Spain and Italy on the south to England and west Germany on the north.

Until additional data are secured it is difficult to decide whether this

family originated in Africa (Stehlin, 1899) or in Asia (Schlosser, 1903).

Sabre Tooths.—Stromer**® has recently traced the history of the ma-

cherodonts in north Africa, Asia and Europe. A form closely similar to

44 STROMER, ERNST: ‘Mitteilungen Uber Wirbeltierreste aus dem Mittelpliocin des

Natrontales (Agypten). 38. Artiodactyla: Bunodontia: Flusspferd.” JZeits. d. Deutsch.

Geolog. Ges., Band 66, Abhandl., Heft 1, pp. 1-83. 1914.

45 STROMER, ERNST: “Mitteilungen Uber Wirbeltierreste aus dem Mittelpliocin des

Natrontales (Agypten).”” Zeitschr. d. Deutsch. Geologisch. Gesellschaft, Band 65, Ab-

handl., Heft 3, pp. 350-372. 1913.

964 ANNALS NEW YORK ACADEMY OF SCIENCES

M. aphanistus Kaup is the first known member of the sub-family to be

discovered in Africa. He observes that the hyenas and otters (Lutra),

the seals (Pristiphoca), and sabre tooths (Macherodus) speak strongly

for a connection between North Africa (Egypt), Asia and Pliocene Eu-

rope. The sabre-tooth survives into the First, Second, and possibly into

the Third Interglacial Stage (see note on Chellean culture).

Moose (Alces).—The earliest representative of the moose*® is the Alces

latifrons Dawkins from the Forest Bed of Cromer. It also occurs in the

sands of Mosbach and of Mauer (Second Interglacial) and in the sands

underlying the lower travertine layers of Taubach (Third Interglacial),

always distinguished by the relatively simple palmation of its antlers

from the existing A. machlis. The animal is also recorded in the Third

Interglacial layer of Rixdorf and it survived in Germany into post-Neo-

lithic times.

4 PROBLEMATIC EVIDENCE OF MAN

Saint-Prest is the most ancient Lower Pleistocene deposit in the basin

of Paris.*7 Coarse sands and gravels form part of the “high terrace”

50m. above the present level of the river Eure and contain a First Inter-

glacial fauna of Hlephas meridionalis and Equus stenonts as well as the

“eoliths” known as Reutelren. This constitutes the Saint-Prestien stage

of Depéret.

The locality of Saimt-Prest is famous because in 1863 Desnoyer first re-

ported the discovery of a number of bones with incision lines which he

considered to be the work of man. These deposits were at the time re-

garded as Pliocene and gave rise to the theory of the occurrence of man

in Plhocene times. The human origin of these incisions has long been a

matter of dispute and is still doubtful. The associated fauna at Saint-

Prest includes the southern elephant, the etruscan rhinoceros, the hippo-

potamus, the giant beaver, three species of beaver and one of the bison.

There is thus little doubt that this deposit is of First Interglacial age.

Supposed confirmation of Desnoyer’s discovery was the alleged finding

by Abbott of several worked flints, two in situ, in the Cromer Forest Bed.

This question has become more or less identified with the eolithic

theory which postulates a long stage of the artificial use of flints antece-

dent to the pre-Chellean and Chellean Stages, which are here considered

as belonging in the Third Glacial Stage, although some authors place the

pre-Chellean in the First Interglacial Stage.

4 DripTRicH, W. O.: “Neue fossile Cervidenreste aus Beeacnane Jabhreshefte des

Vereins f. vaterliindische Naturkunde, Jahrg. 66, pp. 320-336. 1910.

47 HAUG, EMILE: “Traité de Géologie. II. Les Périodes géologiques,” p. 1807. Libr.

Armand Colin, Paris, 1908-1911.

OSBORN. REVIEW OF THE PLEISTOCENE 265

Eolithic Theory.s—Following Desnoyer’s discovery in 1863 was that

of L’Abbé Bourgeois in 186%, who found in the Miocene of Thenay,

Loire-et-Cher, flints supposed to be the work of man. In 1877 Rames

brought to notice flints from, the Upper Miocene volcanic ash beds of

Puy-Cournay, Cantal, in central France. In 1892 Brown proposed the

term “eoliths” to distinguish these supposedly primitive artifacts from

the “paleoliths” of Lubbock (Fig. 13). The Belgian geologist Rutot

has devoted many years to the development of the eolithic theory and has

Fic. 13.—Zolithic, Paleolithic and Neolithic implements

A Holith, Mafflean Epoch, Belgium. B Paleolith, Chellean Epoch, Milton Street,

Kent, England. COC Neolith, Upper Robenhausian Epoch, Gille Leie, Denmark. Photo-

graph by MacCurdy, 1909.

attempted to prove that like the Paleolithic the Holithic period is capable

of subdivision into a number of stages or industries which are geolog-

ically demonstrable.

The supposed eolithic flints are very rough, but rude as they are they

generally exhibit one part shaped as if to be grasped by the hand while

the other part appears to be edged or pointed for cutting.*® It is gener-

48 MacCurpy, G. G.: “The Eolithic Problem. Evidences of a Rude Industry Antedat-

ing the Paleolithic,” Amer. Anthropol., N. S., Vol. VII, No. 3, pp. 425-479. July-Sept.,

1905. ‘

#7PpncK, A.: “The Antiquity of Man.’ Lecture before Washington Acad. of Sci.,

Feb. 1, 1909. Abstr. Science, N. S., Vol. xxix, No. 739, pp. 359-360. Feb. 26, 1909.

266 ANNALS NEW YORK ACADEMY OF SCIENCES

ally admitted that these flints are mostly of accidental shapes and show

little or no proof of being fashioned by-human hands. The chief diffi-

culty in the eolithic theory is that flints of a similar character occur in

deposits of Upper Oligocene age (Rutot) and may even be found in de-

posits of Lower Eocene age (Breuil). The leading French archeolo-

gists, Cartailhac, Breuil, Obermaier, Boule, accordingly reject the

eolithic theory entirely. Breuil considers that the various eolithic shapes

are entirely accidental products due to pressure of closely compacted

flints. Boule observes:°° “As to the ‘eoliths’ I have combatted the the-

ory not only because it seems to me improbable, but because a long

geological experience has shown me that it is often impossible to distin-

guish stones split, cut or ‘retouched’ by purely physical agents from

certain products of rudimentary workmanship.”

It does not appear that the eolithic theory has been strengthened in

recent years. If eoliths are to be regarded as human artifacts the an-

tiquity of man or of a prehuman type given to shaping stone implements

would be vastly increased. It is probable that the ancestors of early

man possessed grasping power of the hand. The only known Miocene

and Pliocene primate of Europe which might be considered as an eolith-

maker or eolith-user is Dryopithecus ; all the other known pre-Pleistocene

primates belong to some one of the existing phyla of monkeys, baboons,

or apes.

Pithecanthropus erectus——Of late Phocene or early Pleistocene age is

the Pithecanthropus erectus of Trinil, Java, discovered by the Dutch

army surgeon Eugen Dubois in the year 1891. The remains were re-

corded by Dubois as of Upper Pliocene age because of their association

with a rich fauna containing Stegodon ganesa, Elephas hysudricus, and

other mammals similar to those of the Upper Pliocene of the Siwaliks of

India. Reéxamination of the NKendeng stratum in which Pithecan-

thropus occurs, by Holz (1907), Elbert (1908) and others appears to

demonstrate that the Pithecanthropus occurs in strata corresponding

with an early Pluvial period following a period of low temperature which

would appear to accord with the early Pleistocene of Europe. It is there-

fore generally agreed that Pithecanthropus is of early Pleistocene age.

Following the monographic description of this very important type by

its discoverer Dubois®? was the keen analysis of Schwalbe,°? who regards

5° BOULE, MARCELLIN: “L’Homme fossile de la Chapelle-aux-Saints.”” Ann. de Paleont.

Vol. VI, pp. 111-172, pll. XVII-XX, 1911; Vol. VII, pp. 21-192, pill. IV-XIx, 1912;

Vol. VIII, pp. 1-71, 1913. ,

= DuBois, Euc.: “Pithecanthropus erectus. Eine Menschenaehnliche Uebergangsform

aus Java.” 4to. Landesdruckerei, 39 pp., 2 pll. Batavia, Java, 1894.

52 SCHWALBE, G.: “‘Ueber fossile Primaten und ihre Bedeutung fiir die Vorgeschichte

* des Menschen.” Mitteil. Philomat. Ges. Elsass-Lothringen, Vol. IV; No. 1, Decade 16.

1908. Strassburg, 1909.

OSBORN, REVIEW OF THE PLEISTOCENE 267

these remains as representing either a direct or indirect ancestor of a

human phylum which stands intermediate between that of the apes and

Neanderthal man (H. neanderthalensis) both in respect to its high brain

capacity and the structure of its femur. ‘The unusually straight femur

indicates an erect attitude, to which the specific name P. erectus applies;

in fact, from the structure of the thigh bone Dubois concluded that the

“Trinil Ape-Man” had free use of the arm and hand, which were now no

longer required for locomotion, and that the hand was already far ad-

vanced in the line of differentiation which developed it into an organ of

touch and capable of fashioning tools.

LIFE OF THE MEDITERRANEAN ISLANDS

It appears that the continental elevation of southern Europe in early

Pleistocene times established migrating routes or land connections be-

tween the islands of the Mediterranean with Europe on the north and

Africa on the south. Riitimeyer (1869) was one of the first to maintain

that north Africa, including Morocco, Algeria and Tunis, was stocked

with mammals by way of Gibraltar and perhaps also by way of Sicily

and Malta. In the islands of Malta, Cyprus and Crete as recently ex-

plored by Bate’* we have proof, first, of a long period of connection with

the neighboring continents through elevation, and second, of the isolation

of the islands through subsidence. The isolation is followed by the

dwarfing of several types of large mammals which, confined on the islands

were made captive by the sea. Pohlig believes** that toward the end of

the First Glacial Stage large mammals migrated to Sicily which at that

time was connected both with Europe and Africa. The land bridges

then became submerged and the large mammals dwindled in proportions

through interbreeding and isolation into dwarf races. It appears prob-

able that Cyprus became isolated as an island first, because the extinct

Elephas cypriotes and Hippopotamus minutus are both more primitive

than the species of Malta and Sicily. That Malta retained its connec-

tion with Sicily for a long period is indicated by the common occurrence

in the cavern deposits on both islands of the two species Elephas mnaid-

rensis and Hippopotamus pentiandi. The dwarf elephant race (E.

melitensis) characteristic of Malta has also been found on the mainland

near Rome, which would seem to indicate that land connection between

the Italian mainland and Malta was renewed more than once.

53 BaTE, D. M. A.: “On Hlephant Remains from Crete, with Description of Elephas

ereticus sp. n.,”? Proc. Zool. Soc. London, pp. 238-250. Aug. 1, 1907.

54 PoHLIG, H.: “Eine Elephantenhoéhle Siciliens und der erste Nachweis des Cranial-

domes von £lephas antiquis,’ Abhand. kinigl. bayer. Akad. Wissensch., el. ii, Bd. xviii,

Abth. 1, pp. 75-108, pll. i-v, 1895. Sep. Munich, 1893.

268 ANNALS NEW YORK ACADEMY OF SCIENCES

The diminutive elephants of the Mediterranean islands were all de-

scendants of the straight-tusked species H. antiquus. 'The researches of

Bate confirm this relationship. They attained a height not exceeding

five feet. The adaptability to which LZ. antiquus owed its wide geographic

distribution and continued existence through a long period of time may

account for its survival in the Mediterranean islands despite rapid dim-

inution in size under adverse circumstances. The true African elephant

(Loxodonta) never crossed the Mediterranean.

The reduced existing fauna of the Island of Cyprus contains a min-

cling of Eurasiatic and north African mammals and shows the effects of

deforestation in historic times. Descendants of Eurasiatic ancestors pre-

vail in the Mediterranean islands. The recently discovered Myotragus

balearicus of the Pleistocene cave deposits of the Island of Majorca is

now regarded as related to the Rupicaprine or Alpine chamois type

(Andrews).

SECOND GLACIAL STAGE—SAXONIAN, MINDEL, KANSAN

The second glaciation was the greatest both in Europe and America.

We observe that the most extended drift sheets are those in the Scandi-

navian region, on the British Isles, around the northern Swiss Alps, and

from the Keewatin center west of Hudson Bay in British America. ‘The

whole rise and fall of the Mindel glaciation in the Alps is estimated by

Penck as occupying a very long period of time. The snow line in the

Alpine region descended 1,300m. lower than at the present time.

The only notable exceptions are in the Labrador region of eastern

North America where the main ice field was formed at a later stage,

known as the Illinoian. It also appears that the Third Glacial or Riss

drift of the western Alps is the greatest in that region and of similar age

to the Illnoian.

In this second glacial advance the Scandinavian ice field reached its

farthest southerly limits. In northwestern Europe this main Saxonian

(Geikie) glaciation extended to the northern slopes of the Carpathians,

the Sudetes, the Erz Gebirge, the Thuringian and the Harz Mountains.

From these ice sheets were given off the “Older Drift,” or “Lower Dilu-

vium” of northern Germany, and in the Swiss Alps this Second glacia-

tion sent out its Mindel drift as the most extensive fringe along the

northern borders of the Alps; on the eastern and southern borders of the

Alps the Second Glaciation was about as extensive as the Third glacia-

tion; on the western borders of the Alps the Second glaciation was less

. extensive than the Third. Similar conditions prevailed in America;

from the Keewatin Centre the ice cap extended its drift southward into

OSBORN, REVIEW OF THE PLEISTOCENE 269

Missouri, Lowa, Kansas and Nebraska beyond the hmits both of the First

and the Fourth glaciation.

Thus the Saxonian drift of North Germany, the Mindel drift of the

Swiss Alps and the Kansas drift of America are correlated (Penck, Lev-

erett) both by their great antiquity and by their very wide extent. The

eminent geologist Wahnschaffe, however, correlates the “Old Drift” of

the north German lowland with the Third or Riss glaciation imstead of

with the Second.

SECOND INTERGLACIAL STAGE—MINDEL-RIss

Penck regards the Second Interglacial or Mindel-Riss Stage as by far

the longest of the interglacial intervals in the Alpine region, estimating

the period between the maximum Second glaciation (Mindel) and maxi-

mum Third (Riss) as high as 360,000 years. In America also by com-

paring the erosion of the Second Glacial (Kansan) drifts with those of

the Third Glacial (Illinoian) drifts it would appear that the Second

Jnterglacial Yarmouth Stage was of greater duration than the entire

interval between the Third Glacial and present time. In the north Ger-

man lowlands it is shown to be a long interval from the amount of sedi-

mentation effected by the interglacial rivers and streams, but whether in

this region it is longer than the First Interglacial Stage is doubtful (Ley-

ereit, 1910, p. 273).

MOISTURE FOLLOWED BY ARIDITY

In course of this long warm Second Interglacial Stage the climate

again moderated, becoming slightly warmer than the climate of to-day.

The climate immediately following the retreat of the ice was cool and

moist, then followed a long warm stage, but this stage was finally suc-

ceeded by a period of aridity both in Europe and America in which the

first loess deposits occurred. In Russia also the Second Interglacial seems

to begin with a cool and moist phase followed by a more arid or steppe-

like climate favorable to the deposition of loess. It would appear that

the height of the interglacial aridity was reached during the deposition

of the loess. The “Older Loess” deposition certainly began both in Europe

and America during the Second Interglacial Stage although in neither

country is the “Older Loess” so continuous or so thick a deposit as the

“Newer Loess.” In Europe the “Old Loess” lies between the “Old Drift”

of the First and Second Glacial advances and the “Middle Drift” of the

Third Glacial advance. At Mauer near Heidelberg the loess lies imme-

diately above the Upper Mauer sand layer which contains an arctic-

tundra fauna (Forster, 1913). The various layers of loess are of the

270 ANNALS NEW YORK ACADEMY OF SCIENCES

utmost importance both in Europe and America in the correlation of

human and mammalian life, also in their significance as to the climate

of interglacial times. Loess consists of a fine, porous, silicious, calcareous

silt, usually of light brown color, characterized by a peculiar competency

to stand in vertical walls during erosion. Its origin and transportation

are believed to have been partly sub-glacial, partly fluviatile, partly eolian.

The fine mud carried by the sub-glacial streams in glacial times became

desiccated and redistributed by the wind. Penck (1904) describes the

Pleistocene loess as formed in districts traversed periodically by great

streams leaving dry mud which in arid periods was redistributed by

eolian agencies. Its Pleistocene distribution is quite independent of alti-

tude since it occurs in the interglacial deposits of Europe from sea level

to a height of 1,500m.

CLIMATE

A considerable part of the elevation of the Swiss Alps apparently took

place (Penck, 1910) during the Second Interglacial Stage, and this in-

creased altitude is considered by some European authorities to be the

cause of the greater extent of the Third glaciation in the western Alps.

The Hottinger breccia near Innsbruck is referred by Penck (1909, p.

1157) to the Second Interglacial Stage with its rich flora indicating a

climate warmer than that of present times; this breccia hes on one of the

old “High Terraces” of Second Interglaciai times. The plants include

the fir (Pinus sylvestris), spruce (Picea sp.), maple (Acer pseudopla-

tanus), buckthorn (Rhamnus frangula), several willows (Salix nigricans,

S. glabra, S. incana, S. triandra), the wayfaring tree (Viburnum lan-

tana), yew (Taxus baccata), elm (Ulmus campestris), strawberry (Fra-

garia vesca), self-heal (Prunella vulgaris), beech (Fagus silvatica), and

mountain ash (Sorbus aucuparia), buckthorn (Rhamnus Hettingensis),

related most closely to R. latifolia, now living in the Canary Isles, the

box (Buxus sempervirens), also a southern species; and most important

of all a rhododendron (R. ponticum) which now lives in the Caucasus

five degrees south of the latitude of Innsbruck and in a climate on the

average 3° C. warmer. ‘Taking all the facts into consideration Penck

concludes that the climate of Innsbruck in the days of the Hotting brec-

cia was 2° C. higher than it is now. In correspondence with this the

snow-line stood 1,000 ft. above its present level, and the Alps save for the

higher peaks were almost completely denuded of ice and snow.*°

A picture of the flora of the Second long warm Interglacial Stage is

also afforded in the Quaternary tuffs of Provence, where the remains of

5 SoLtas, W. J.: Ancient Hunters and their Modern Representatives, p. 27. 8vo.

MacMillan & Co. London, 1911.

OSBORN, REVIEW OF THE PLEISTOCENE Aa |

plants are associated with elephants of the /. antiquus stage. “The flora

of the Quaternary tufts,’ observes Saporta,°*® “is composed almost entirely

of woody forms living in valleys and by the sides of streams.” It is for

the most part analogous with the present flora of Provence. Of the

thirty-seven species, twenty-nine still occur in this region. Among the

forms which have since retreated to the south are the sweet bay (Laurus

nobilis) and another species of laurel (L. canariensis) which is now con-

fined to the Canaries. The greater humidity of the time is indicated by

the presence of species of pine which require more moisture. As in the

Norfolk Interglacial, the figs (Ficus) and the Judas trees (Cercis) flour-

ished. The ash (Frazimus) is of a species now found in Corsica and

Italy. On the whole, the forest trees and forest ground flora are surpris-

ingly modern, including oaks, elms, poplars, willows, lindens, maples,

sumacs, dogwood, hawthorn. Among the climbing plants are the vine

(Vitis) and clematis (Clematis).

MAMMALS

This hfe period was first observed by Lyell and Evans in Essex, Eng-

land, and was subsequently recognized in Germany and France. Geo-

logically the deposits are partly of fluviatile origin, consisting chiefly of

river sands and gravels in which the remains of hippopotamus, elephants

and rhinoceroses occur. These animals were formerly cited as proof of

an almost tropical climate, but the evidence of the flora, enumerated

above, and the equally numerous hardy types of animals tend to modify

the former theories as to extremely warm Second Interglacial tempera-

‘tures. The geographic connections of Europe with the south through the

Jand bridges of Lower Pleistocene times still persisted in Italy in whole

or in part, because the depression of the southern portion of the continent

of Europe had not yet begun.

Survivals-—The mammals occurring in these Older Diluvial sands and

gravels include several Pliocene survivals from the First Interglacial

Stage, associated with the etruscan rhinoceros (Dicerorhinus etruscus).

If Montmaurin belongs to this stage we may include Macherodus. At

Mauer two primitive types of bear, Ursus arvernensis and U. deningert,

are recorded, also Trogontherium cuviert. The Mauer horse first identi-

fied as H. stenonis is now referred to H. mauerensis.

Among the chief localities where the river deposits containing the

mammals referred to the Second Interglacial Stage occur are the fol-

lowing :

56 Drm Saporta, G.: ‘La Flore des Tufs Quaternaires en Provence.” C. R. Sess. Congr.

Sci. France. Aix, 1867.

272 ANNALS NEW YORK ACADEMY OF SCIENCES

EARLY PHASE. WARM FAUNA.

Mauer Sands (Lower). near Heidelberg. Germany. Warm Fauna.

Montmaurin (Haute Garonne), Pyrenees, France.

St. Roche. France.

MIDDLE PHASE,

Mosbach, near the Neckar in northern Baden (Fig. 9, 12).

Siissenborn, near Weimar, Germany (11).

LATER PHASE. COLD TUNDRA FAUNA.

Mauer Sands (Upper). near Heidelberg. Germany (14). Cold Fauna

approaching the Third Glacial Stage.

Extinctions—The mammals of this grand life zone have lost nearly

all resemblance to those of Upper Pliocene times with the exception of

the survival of the etruscan rhinoceros and possibly of the sabre-tooth

tiger. The polycladine deer of Upper Pliocene times and of the Norfolk

Forest Bed, or First Interglacial, have vanished; neither are there any

traces of the axis deer (C. azis).

Arrivals ——The Second Interglacial Stage is readily distinguished both

in France and Great Britain by a number of important new arrivals,

chief among which are the “old elephant” (#. antiquus) and the broad-

nosed rhinoceros (D. merckiu). Another very important arrival is the

hon related to the African Felis leo. The southern elephant has now

certainly passed into the Hlephas trogontheru stage of Pohlig for the

type specimen of this intermediate species occurs at Siissenborn; in fact,

this is the “H. trogontheru stage” of Pohlig: it is also known as the

“older FE. antiquus” stage by Schmidt and other authors. The southern

mammoth F£. (meridionalis) trogontherwt is replaced by the more pro-

gressive and typical #. trogontherit. The broad-faced moose (Alces latt-

frons), the giant deer (Megaceros) and the roe deer (Capreolus) are all

present in the cooler and forested phases of this interglacial period. The

true stag (Cervus elaphus) is certainly recorded. The cattle (Bos primi-

genius) begin to be numerous and the bison (Bison priscus) also appear

in numbers. Horses of larger size occur (EZ. mosbachensis, EH. stssen-

bornensis).

Among the river-living forms are the beavers (Castor). The giant

beaver (T'rogontherium) is by some authors said to make its last appear-

ance in Europe in this sub-stage, but it is again recorded in the Third

Interglacial at Chelles. Other rodents include the marmots (Marmotta)

now found in the Alps, Carpathians and Pyrenees, whose remains may

have been borne down by the streams. Beside the lions the carnivores -

include the typical Eurasiatic forest forms, namely, the lynx (F. lynz),

two varieties of bear (U. deningeri, U. arvernensis), and the badger

(Meles).

The chief components of the fauna of the Second Interglacial Stage

OSBORN, REVIEW OF THE PLEISTOCENE 273

are seen to belong, first, to the Eurasiatic Forest and Meadow Fauna,

only separated by specific and sub-specific differences from the Prehis-

toric Fauna of Europe; second, to the surviving African-Asiatic fauna,

including the hippopotamus, two very distinct kinds of elephant, and two

rhinoceroses; third, there is evidence in the late colder phases of this

period of the first occurrence in Europe of the Tundra Fauna as repre-

sented by the reindeer (Rangifer tarandus). This animal is recorded in

the gravels of Siissenborn by Weiss. Huilzheimer®’ also speaks of the re-

mains of reindeer as occurring both in Stissenborn and Steinheim in asso-

ciation with the remains of L. trogontheru. This author regards L. tro-

gontherw from the structure of its grinding teeth as analogous in habit

to the Asiatic elephant which inhabits the forests of India, and believes

that the presence of this animal indicates a relatively moist climate and

well forested country.

: In this assemblage it is noteworthy

ee re TYPES that the Eurasiatic Forest and

Primates ;

Homo heidelbergensis hardy temperate types greatly pre-

ee keer AGW A dominate over the African-Asiatic

Straight-tusked elephant types. This is another indication

E. antiquus that the climate was of a warm-tem-

Trogontherian mammoth

E. trogontherii

Broad-nosed rhinoceros

perate character rather than such as

now characterizes southern Asia and

D. merckii Africa. It follows that all the Afri-

Etruscan rhinoceros can-Asiatic mammals may have been

D. etruscus well protected by hairy covering and

Hippopotamus

adapted to a temperate climate.

H. major :

Be Sabretooth tizer In the caverns near Montmaurin

Macherodus in the Pyrenees®® we find remains of

Lion an early Pleistocene fauna which

Felis leo spclea contains the sabre-tooth tiger (MM.

EURASIATIC Harpy FAUNA

Urus

Bos primigenius

latidens), the broad-nosed rhinoc-

eros (D. merckiw), the stag (C. ela-

Bison phus), the brown hyena (H. brun-

Bison priscus nea striata).

Stag. roe deer, moose, giant deer The most typical fauna is that of

fe ee eae Mosbach in northern Baden. Here

e : wy

deer there occur all the characteristic

Rangifer tarandus mammalian types of the period, the

hippopotamus, the urus, the bison,

5? HILZHEIMER, MAx: Handbuch der Biologie der Wirbeltiere, pp. 678-679. Stuttgart,

1912-1913.

58 BouLn, M.: “La Caverne & Ossements de Montmaurin (Haute-Garonne).” L’An-

thropol., Vol. XIII, pp. 305-319. 1902.

44 ANNALS NEW YORK ACADEMY OF SCIENCES

the broad-nosed rhinoceros, two species of mammoth (F. antiquus, E.

trogontherw), and the horse (#. mosbachensis) .

In the Lower sands of Mauer near Heidelberg there occur the first re-

corded remains of man and a fauna including some primitive species.

Homo heidelbergensis.—To the faunal stage of Hlephas antiquus and

the etruscan rhinoceros (D. etruscus) is to be added the Heidelberg man,

determined from a lower jaw discovered by Otto Schotensack*? im 1907

in the Lower Mauer Sands at a depth of 24.10 m., one of the most im-

portant discoveries in the whole history of anthropology. The lower jaw

is exceptionally massive, without chin projection, with a large but essen-

Fie. 14.—Sand pit at Mauer near Heidelberg

The lower jaw (Homo heidelbergensis) was found at the spot marked with a cross.

After Schoetensack and MacCurdy.

tially human set of teeth; in other words, it is a jaw in some respects

similar to that of an anthropoid ape but containing the dentition of a

man, namely, typically human canine and molar teeth. The jaw is now

regarded by anatomists as resembling on a very massive and primitive

scale the jaw of the neanderthaloid human type (Homo neanderthalen-

sis) which first occurs in the Third Interglacial Stage.

The fauna associated with Homo heidelbergensis is of an ancient char-

59 SCHOTENSACK, OTTO: Der Unterkiefer des Homo heidelbergensis aus den Sanden

von Mauer bei Heidelberg: Ein Beitrag zur Paliontologie des Menschen. Verlag von

Wilhelm Engelmann. Leipzig, 1908.

OSBORN. REVIEW OF THE PLEISTOCENE Os

acter. Schdtensack likened it to that of the First Interglacial or Nor-

folkian Stage. The presence of the etruscan rhinoceros would appear to

justify this opinion, but it is overborne by the similarity to the fauna of

Mosbach including the presence of Equus mosbachensis, a species highly

characteristic of the Second Interglacial Stage. The entire fauna of

these Lower Sands of Mauer is now identified (Schmidt, 1912), as fol-

lows: Elephas antiquus, D. (Rhinoceros) etruscus,°° Equus mosbachen-

sis, Sus scrofa fere, Alces latifrons, Cervus elaphus, Capreolus capreolus,

Bison priscus, Bos primigenius, Ursus arvernensis, U. deningert, Felts

leo, Felis catus, Canis neschersensis, Castor fiber. The enumeration of

Fic. 15.—Heidellerg jaw

The human lower jaw (%4 natural size) found near Heidelberg, on which is based the

species Homo heidelbergensis. After Schoetensack.

this fauna is very important as indicating the temperate climatic condi-

tions which surrounded the Heidelberg man. Wurm observes that the

Etruscan rhinoceros only occurs in Mauer and that its variations indi-

cate a transition towards the D. merckiw which occurs at Mosbach and

Siissenborn but not in Mauer. Above this layer occurs a deposit of the

“Older Loess,” indicating an arid climate. The Upper Sands of Mauer

contain a cold fauna which by some is referred to the close of the Second

6 WourM, A.: ‘“‘ttber Rhinoceros etruscus Fale. von Mauer a. d. Elsenz (bei Heidel-

berg).”’ Verh. d. Naturhist.-Medizin. Vereins zu Heidelberg, N. F. XII Band, 1 Heft,

pp. 1-62, pll. I-IV. 1912.

Elsenz (bei Heidelberg), I. Felis leo fossilis. Jahresberichte und Mitteilungen des Ober-

rhein. geol. Vereins, N. F., Bd. II, Heft 1, pp. 77-102. 1912.

276 ANNALS NEW YORK ACADEMY OF SCIENCES

Interglacial stage, by others (Wurm,*? 1913, p. 68) to the “Younger

Loess,” that is, to the Fourth Glacial or Postglacial Stage.

CHELLEAN CULTURE WITH ANCIENT INTERGLACIAL FAUNA

In favor of the antiquity of the Chellean culture may be urged the fact

of its association in several localities (Torralba, Abbeville, Piltdown)

with the primitive mammals identified as Macherodus, D. etruscus,

Equus stenonis. ‘The specific identifications may be incorrect, but these

Pliocene species are characteristic of the Second Interglacial Stage and

are not certainly recorded in the Third Interglacial Stage of northern

Kurope at least. For example, at Torralba, Province of Soria, Spain,

there has been discovered (Harlé, 1910, p. 75) an old typical Chellean

camping site containing abundant remains of D. mercku and FE. merid-

tonalis (trogontherit) mingled with remains of other mammals of prim-

itive type identified as Dicerorhinus etruscus and Hquus stenonis. These

associations with Chellean remains tend to support the theory that the

Chellean culture began during the Second Interglacial Stage. Another

very ancient fauna associated with very primitive Chellean or pre-Chel-

lean implements is that found near Abbeville, Gisement de Champ de

Mars.°* Beside typical members (such as H. antiquus, LH. meridionalis

trogontheru, and D. mercku) of warm Second Interglacial times this

fauna is said to contain such primitive types as Trogontherium, D.

etruscus, Equus stenonis, also very numerous specimens of Macherodus

and Hyena brevirostris. |

We cannot fully agree with Schmidt (1912) when he observes that the

faunal separation of the Acheulean and Chellean is not so marked that

we are obliged to separate these cultures by a long period of time.

FAUNA OF THE PYRENEES, CANTABRIAN ALPS, SPAIN AND PORTUGAL *

The entire warm fauna characteristic of Germany, Great Britain and

France also penetrated the Cantabrian Alps, Spain and Portugal as far

south as Gibraltar. ;

A macaque (MJacacus) related to the Algerian species occurs in the

erotto of Montsauné (Haute Garonne) associated with the hyena (4.

62 WurM, A.: “itber eine Neuentdeckte Steppenfauna von Mauer an der Elsenz (bei

Heidelberg).’’ Jahresber. u. Mitt. d. Oberrhein. geol. Vereins, N. F., Bd. III, Heft 1,

pp. 62-78, pl. vi. 1918.

6&8 P’AULT DU MESNIL, G.: “Note sur le Terrain Quaternaire des Environs d’Abbe-

ville.’ Revue Mensuelle de l’&cole d’Anthropologie de Paris, VI year, pp. 285-296,

1896. '

6 HARLE, Epovarp: ‘“‘Les mammiféres et oiseaux quaternaires connus jusqu’ici en

Portugal. Memoires suivi d’une liste générale de ceux de la Peninsule Ibérique.” Com-

mun. du Service Géol. du Portugal, T. viii, pp. 22-85, pll. I-V. 1910.

OSBORN, REVIEW OF THE PLEISTOCENE OV"

striata), with a dhole (Cyon), and other members of the warm fauna of

EH. antiquus and D. mercku. The porcupine (Hystriz cristata) also

occurs here. 7

The striped hyena (H. striata brunnea) is associated with Machero-

dus latidens in the cavern of Montmaurin (Haute Garonne). The

striped hyeena occurs at five other localities in the Pyrenees, Spain and

Portugal (Furninha) ; it has also been recognized in Germany (Mos-

bach), Austria (Hundsheim) and France (Harlé, 1910, p. 40); it dis-

appears later or retires to the south, while the spotted hyzna (H. cro-

cuta spelea) becomes adapted to the extreme cold and survives with the

reindeer to the end of Postglacial times. The Carnivora of this region

are Felis leo spelea, the panther (Felis pardus), the wild cat (Felis

catus), and the lynx (Felis pardina).

SECOND AND THIRD GLACIAL AND INTERGLACIAL EPpocus

AFRICAN-ASIATIC FAUNA

Four great animals especially characterize this fauna: Hlephas trogon-

theru. and Hippopotamus major, Dicerorhinus mercku and Elephas an-

tiquus.

Old Elephant (Hlephas antiquus).*°—The “old elephant” or straight-

tusked elephant (Hlephas antiquus) does not occur in France or Great

Britain until the Second Interglacial Stage, but it is said to occur in the

Arno valley of Italy during an earlier stage in which it is associated with

a warm fauna including the southern mammoth and the hippopotamus.

The typical H#. antiquus is recognized by its narrow, elongated grinding

teeth with comparatively few plates which, combined with its skull char-

acters, suggest its affinity to the modern African (Lozodonta) rather

than to the Indian elephant (Huelephas) group. While during the first,

or Norfolk, interglacial period it is confined to Italy, in subsequent in-

terglacial times it wandered into northern Europe as one of the grandest

and most distinctive forms, attaining a very wide distribution. Pohlig

certainly overestimates its size®* in assigning to it a height of 5 m. at the

back (16 ft. 8 in.), or 1m. more than the mammoth, and with tusks also

5m. in length. In consequence of the size and weight of the tusks, the

head, shoulders and fore legs were enormously developed. The same

writer believes that the habitat of.this mammoth retreated and advanced

% Ponuic, H.: “Dentition und Kraniologie des Elephas antiquus Fale. mit Beitrigen

liber Hlephas primigenius Blum. und EHlephas meridionalis Nesti.” Noy. Act. Ksl. Leop.-

Carol. Deutsch. Akad. Naturforsch., Vol. LIII, No. 1, p. 326. Halle, 1888,

6 POoHLIG, H.: Hiszeit und Urgeschichte des Menschen. Leipzig, 1907.

ANNALS NEW YORK ACADEMY OF SCIENCES

AN)

with the successive ice waves and warm interglacial times. Because of

the resemblance of the grinding teeth of HE. antiguus to those of the

African elephant (L. africanus) it has been assumed perhaps too readily

that this ancient elephant was characieristic of a tropical climate. It

resembles the African elephant in the prominence of the enamel bands

of the grinding teeth, which are adapted to the comminution of twigs

and woody food, which justifies the belief that this animal frequented

the forests. For these reasons Hilzheimer regards L. antiqguus as indica-

tive of forest conditions. ;

Rhinoceroses—The three great rhimoceroses characteristic of the Eu-

ropean Pleistocene are each of distinct geological value. Im general

D. etruscus belongs to the First Interglacial Stage, D. mercku character-

izes the Second Interglacial Stage and most of the Third Interglacial,

while D. antiguttatis is distinctive of the Fourth Glacial and the Post-

glacial.

The two species first named are apparently related to the Sumatran

phylum (Dicerorhinus sumatrensis). The D. eiruscus of the Val

d’Arno, of the First Interglacial and of the early phases of the Second

Interglacial is a relatively small animal, distinguished by brachyodont

grinding teeth and long, slender limbs, a small anterior and a larger

posterior horn. It is remotely related to the Sumatran rhinoceros but

differs in the absence of cutting, or incisor teeth. It is essentially a

browsing type. Its remains in Mauer are said (Wurm, 1912) to afford a

transition to D. merckit.

Related to this animal in the Second Imierglacial Stage in Great

Britain, Germany, France, Italy, there appears the broad-nosed rhimoce-

ros known as D. megarhinus, or D. mercku. It resembles D. etruscus in

its smaller anterior and larger posterior horn and im the elongation of

its limbs and feet, but differs from it in the possession of relatively long-

crowned (hypsodont) grinding teeth adapted to grazing habits. This

animal is very widely distributed geographically in the Second and the

first half of the Third Interglacial Stage, and is in most localities asso-

ciated with remains of the hippopotamus and “old elephant.”

Quite distinct from these animals is the woolly rhinoceros (Diceros

antiquitatis, D. tichorhinus) which belongs with the colder climates of

tundra and steppe conditions and is almost invariably associated with

remains of the true woolly mammoth (E£. primigenius). Like the above

described Sumatran species it lacks the front, or cutting teeth and has in

consequence been improperly considered as related to Dicerorhinus, but

really belongs to the modern African group of Diceros, resembling espe-

cially the species D. simus, with which it closely agrees in its dolicho-

OSBORN, REVIEW OF THE PLEISTOCENE 29

cephalic cranial proportions, its long-crowned teeth, and especially in the

presence of a square upper lip and very large anterior horn and small

posterior horn. It is thus distinguished both by the proportions of its

horns and by the characters of its teeth and lips from the two species of

Dicerorhinus. It is distinctively a grazing animal.

SYNONOMY OF RHINOCEROS GENERA

1. Indian rhinoceros: R. indicus — RHINOCEROS Linnzeus 1758

2. Black rhinoceros of Africa: D. bicornis — DicERos Gray 1821

& Woolly rhinoceros: R. antiquitatis (Blumenbach 1799) — C@Loponta Bronn

1831

4. Sumatran rhinoceros: D. swmatrensis — DICERORHINUS Gloger 1841

5. White rhinoceros of Africa: D. simus = CERATORHINUS Gray 1867

The names of these three rhinoceroses are almost hopelessly confused

in the early literature. The animals converge toward each other in sev-

eral characters, namely, in the loss of cutting teeth and in the develop-

ment of an osseous septum for the support of the nasal bones. The

woolly rhinoceros (D. antiquitatis) is first recorded in Europe in deposits

correlated with the Third Glacial or Riss Stage, the Mammut Lehm of

Cannstatt (Koken, Schmidt, 1912).

Bovines.—The bison (B. priscus) rivalled the mammoth as a wanderer

and was able to adapt itself to wide diversities of climate in Europe.

Asia and America. Originally of African-Asiatic origin it became thor-

oughly acclimated as a Eurasiatic meadow and plains type and may have

extended also into the forests like the existing woodland bison (B. atha-

basce) of Canada. It is readily distinguished as brachycephalic while

its contemporary, the gigantic urus, is long-headed (dolichocephalic),

as well as less agile than the bison. In external appearance, as depicted

in the very numerous engravings and paintings in the Font de Gaume

and other caverns, this animal resembled the existing American bison

(B. americanus) more than the still surviving Lithuanian and Caucasian

form (B. bonasus). The animal appears in the First Interglacial, or

Norfolkian Stage in France. In the Second long warm Interglacial

Stage there existed a bison (B. priscus antiquus) which enjoyed a wide

distribution. The animal found its way to the Mediterranean islands

and gave rise to the pigmy varieties.

The wild ox (Bos primigenius) also occurs in the First Interglacia!

Stage and survived the vicissitudes of the entire Pleistocene Epoch. The

“arus’”’ of Ceesar survived in its wild state in Europe as late as the seven-

teenth century A. D., where it was still to be found in the forests of

Poland and in a few game preserves. It then disappeared so completely

980 ANNALS NEW YORK ACADEMY OF SCIENCES

that even its popular designation “‘auerochs” was transferred to the

Lithuanian bison.®*’ The designations of these two types are therefore

very confusing and are distinguished by Kobelt as follows:

Urus (Pliny), wild ox, urochs, auerochs (Old Ger-

man), tur (Polish), urstier — Bos primigenius

Bonasus (Aristotle) wisent or wisont, subr (Polish), auer-

ochs (of recent date) — Bos priscus

The relations of the wild cattle of Asia to domestication will be con-

sidered on a later page.

Megaceros giganteus——During the first half of the Pleistocene this

noble animal was widely distributed in Ireland, England, Scotland, the

Isle of Man, France, Denmark, Germany, Austria, northern Italy and

parts of Hurasia even into Siberia. The famous Megaceros beds of Ire-

land are freshwater clays which frequently underly the peat bogs. As

observed by Williams these are “boulder-clays” which were redistributed

as lake sediments and accumulated under genial or temperate climatic

conditions lke the present. Owing to the similarity in the palmation

of its antlers the giant deer has been generally (Lydekker, Weber,

Trouessart) placed within or very close to the subgenus Dama, the

fallow deer; but Lonnberg*®* regards the likeness between the giant deer

and fallow deer as convergence and considers that the giant deer is more

closely related phylogenetically to the reindeer, but it is nevertheless so

specialized as to hold an independent place in the system of Cervide.

The Megaceros last appears in early Postglacial times associated with

the Aurignacean culture in Germany; it is not recorded (Schmidt) with

the succeeding Solutrean or Magdalenian culture. It thus became ex-

tinct before the close of the Paleolithic.

Hlaphine or Red Deer.°°—Sir Victor Buck held that the Cervide orig-

inated in Asia and from there spread westward into Europe or eastward

into America. The Asiatic origin of the red deer race has since been

ably maintained by Koppen. A very large race of late Pleistocene times

has been compared by Nehring with the C. canadensis of North America.

Reindeer.—The reindeer of Pleistocene times are generally referred to

the Barren Ground or Tundra type. In this type, which is typified by

the existing Old World reindeer (FR. tarandus, R. spitzbergensis) and by

the American arctic forms (R. arcticus, R. Grenlandicus, R. grant, RB.

8’ POHLIG, H.: Hiszeit und Urgeschichte des Menschen, p. 131. Leipzig. 1907.

RUTIMEYER, L.: “‘Die Fauna der Pfahlbauten der Schweiz.’”? Neue Denkschr. schweiz.

Ges. gesam. Naturwiss., Vol. xix, pp. 68-112. Zitirich, 1862.

KoBre.Lt, W.: Die Verbreitung der Tierwelt. S8vo. Leipzig, 1902.

88 LONNBERG, E.: “‘Which is the Taxonomic Position of the Irish Giant Deer and Allied

Races?” Ark. Zo6l., Vol. 3, No. 14, pp. 1-8. Upsala, 1906.

68 Scuarrr, R. F.: The History of the European Fauna, pp. 246-251. London, 1899.

OSBORN, REVIEW OF THE PLEISTOCENE IS1

pearyt), the antlers are round, slender and long in proportion to the

relatively small size of the animal, while the spreading beam and brow

tines are as a rule but little palmated, although in some forms the brow

tine is palmated.

The woodland type, which is now extinct in Kurope, is typified by sev-

eral American species’? (ft. caribou, R. montanus, Rk. osborni) in which

the antlers are heavier, flatter, thicker, and more heavily palmated on

the spreading beam and on the “brow tine” especially, while the tine

above the brow, which corresponds to the bez-tine of the stag (Cervus),

is elaborately developed and palmated thus contrasting sharply with the

simple bez-tine of the Barren Ground group.

Some writers’! (Scharff) maintain that the Barren Ground reindeer

entered Europe first during the First and Second Glacial Stages while

the woodland group first appears in the Third Glacial Stage. Others

(Hilzheimer) maintain that all the known Pleistocene reindeer belong

to the Tundra form and not to the woodland form. Again, Dietrich™?

recognizes a woodland caribou in the “high terrace” gravels of Steinheim

in the valley of the Murr.

Carnwores.—The larger Pleistocene carnivores embrace the wolves:

(Canis lupus), the bears (U. arctos, U. spelea), the hyenas (H. crocuta

spelea, H. (brunnea) striata), the leopards (Felis pardus of Spain)

and the lions.

The chief enemies of the wild horses and cattle of the Pleistocene were

the lions (Felts leo spelea), descended either from the great cats of the

Pliocene of France and Italy (Felis arvernensis) or more probably mi-

grants from northern Africa. These lions are known from deposits in

England, Belgium, Austria, southern Russia, France, Spain,” Italy,

Sicily, Greece and Algeria."* The fact that remains of this animal are

so often associated with those of the cold Postglacial fauna makes Neh-

ring’s’ suggestion seem plausible that the cave lion was a northern race

of the recent African and western Asiatic lion adapted to a colder cli-

mate and with a heavy coat. After examination of specimens from cen-

tral and northern Europe Boule™* reaches the conclusion that these lions

7 GRANT, Mapison: ‘‘The Caribou.” Ann. Rept. N. Y. Zoél. Soc., no. 7, pp. 175-196.

New York, 1892.

POD Clt., «p> Lb.

7 DietTrRicH, W. O.: “Neue fossile Cervidenreste aus Schwaben,” Jahreshefte des

Vereins f. vaterliindische Naturkunde, Jahrg. 66, pp. 320-336. 1910.

3 HARLE, Epovarp: ‘Les mammiféres et oiseaux quaternaires connus jusaiiei en

Portugal. Memoires suivi d’une liste générale de ceux de la Péninsule Ibérique.’’ Com-

mun. du Service géol. du Portugal, T. VIII, pp. 22-85, pll. I-V. 1910.

7 BouLE, MARCELLIN: ‘‘Les Grands Chats des Cavernes.” Ann. de Paléont., Vol. I, pp.

20-27. Jan., 1906. .

* NEMRING, A.: Uber Tundren und Stenpen der Jetzt-und Vorzeit. mit besonderer

Berticksichtigung ihrer Fauna. Berlin, 1890.

289 ANNALS NEW YORK ACADEMY OF SCIENCES

are not related to the tiger (F. tigris) as was supposed formerly by De

lainville and Lartet. While rich in individual variations Felis leo

spelwa is nearer the lion than the tiger in most of its characters; it

should, in fact. be considered a veritable race of the recent lion with the

name Felis leo spelea. Jt differs from both the recent lion and tiger in

the more gentle and uniform slope of its facial profile and in its large,

fiat forehead, but its limb bones are longer and proportionately thicker.

It sometimes equals and often surpasses the existing lions and tigers in

size. It is represented in the cave engravings and drawings both of

early and late Postglacial times.

The cave hyena (Hyena crocuta spelea) is a variety of the hving

spotted hyzna (Hyena crocuia) of East Africa, but it attamed dimensions

considerably greater than that of its living ally. Ii has the larger propor-

tions, the heavier build, the broad skull, the powerful carnassial teeth

which distinguish the spotted from the striped hyena (H. striata) of

the present day. Although proportionately heavier the hind limbs may

have been shorter than in the spotted hyena in adaptation to the cavern

life which the inclement climate made necessary. The cave hyena was

a very abundant type and is responsible for scattering of the vast num-

bers of the bones of the contemporary animals in a manner not pleasmg

to the palzontologist.

In the caves of southern France a variety (Hyena priscus) of the

striped hyena (Hyena striata) also occurs and there are also discovered

here additional remains of an animal (H. intermedia) resembling the

cave hyena. Thus the Pleistocene species of European hyenas under-

went an evolution of their own. As a result the living African forms

differ more from the Pleistocene hyenas of Europe than they do from

those of Phocene times.

Harlé** records the striped hyena (H. striata) as characteristic of the

earlier or warm Pleistocene of Spain and Portugal; the cave hyena (H.

crocuta spelea) survived into the late Pleistocene through adaptation to

the cold climate. ;

The cave wolf (Canis lupus spelea), a member of the forest fauna,

also attained dimensions greater than its living allies. According to

Gaudry and Boule,“* (1892) no constant osteological differences can be

determined between the Pleistocene cave wolf and the modern wolf of

western Europe, although the cave form is of- considerably larger size.

This animal is represented in the Upper Magdalenian paintings of Font

de Gaume.

7 Op. cit., 1910. pp. 46, 70.

“= Gaupey, A.. and Bovunte, M.: Matériaux pour l’'Histoire des Temps Quaternaires,

Fase. 4. Les Oublietites de Gargas. pp. 108-112. Paris. 1892.

OSBORN, REVIEW OF THE PLEISTOCENE 223

THIRD GLACIAL STAGE—ILLINOIAN, POLANDIAN, RIss

In North America the Third Glacial Stage is heralded by the advance

of a great ice cap radiating from Labrador which sent its glaciers to the

south and far southwest, depositing the Illinoian drift which is regarded

(Leverett, 1910, p. 315) as of an earlier period than the Polandian or

“Middle Drift” of northern Germany or the Riss drift of the Alpine

region. This Third glaciation of the Alpine region has a period of ad-

vance and retreat which is relatively estimated by Penck at 20,000 years,

the snow line descending 1,250m. With it are associated the “high ter-

race” deposits of the Alpine region. The Third glaciation was greatly

extended along the Rhine, in parts of Switzerland, in France, and in

the valley of the Po (Fig. 9). In northern Germany the principal rea-

son for separating the “Middle Drift” (Polandian) from the “Upper

Drift” (Mecklenburgian) is the presence of loess deposits between them

which seems to strengthen the evidence for a Third Interglacial interval.

‘These loess deposits are regarded by certain German geologists (Koken)

as the continuation of the “Older Loess” but by Penck and Leverett they

have been regarded as belonging to the “Newer Loess.”

THIRD GLACIAL FAUNA

The recurrence of a cold climate in Germany is heralded in the

Upper Sands of Mauer by the arrival of the reindeer and other arctic

types. In the Mammut Lehm of Cannstatt is found a fauna which is

regarded by Koken and Schmidt (1912, op. cit.) as contemporaneous

with the Third Glacial advance. It is noteworthy as containing two

new arrivals from the tundras of the north, namely, the woolly mam-

moth (H. primigentus) and woolly rhinoceros (D. antiquitatis), as well

as the reindeer (f. tarandus). The other members of this fauna in-

clude two species of horse, the giant deer, the stag, the bison and the

urus. “Cannstatt,” observes Schmidt (1912, p. 270), “affords a geologi-

eal and final connecting link between the Second Interglacial fauna of

Mauer and the fauna of Early Paleolithic [or Third Interglacial]

times.” If this fauna actually entered Germany during the cold period

of the Third glaciation it returned to the north with the approach of the

warm-temperate climate of the Third Interglacial Stage, because no

trace of it is found until near the close of the Third Interglacial Stage.

THIRD INTERGLACIAL STAGE—RIss-WURM, SANGAMON

The Third Interglacial Stage is shorter than the Second, its geologic

and faunal characters are more fully known, and it embraces the first

284 ANNALS NEW YORK ACADEMY OF SCIENCES

undoubted remains of human stone industry in Europe as well as abun-

dant remains of man. In the Alpine region this Riss-Wiirm Stage is

indicated by “high terraces,’ which rise 25 to 50m. above the existing

streams. The Riss-Wiirm interval is evidently shorter than that be-

tween the Second and Third glaciation. Penck considers that the depo-

sitions of “Newer Loess” which occurred in the Alps near the close of

- Riss-Wiirm Interglacial times represent a cold stage, since the fauna

which it contains is of the Tundra-Alpine type and the Paleolithic im-

plements found in it are closely similar in workmanship to those found

in deposits subsequent to the Wurm glaciation (Penck, 1909, p. 1159).

He regards this as the “Newer Loess” that was laid down prior to the

Fourth glaciation. Koken and Schmidt, on the other hand (1912),

regard the “Newer Loess” as partly or wholly Postglacial, that is, as

occurring after the Wirm maximum. The Third Interglacial loess of

northwestern Europe was comparatively scanty and discontinuous, from

1 to 5m. in thickness, and contains a terrestrial molluscan fauna as in

America. All the indications are that this loess was deposited by pre-

vailing westerly winds. Also along the Danube the loess is chiefly due

to westerly winds. Penck attributes the scarcity of loess in the southern,

eastern and western borders of the Alps to the presence of thick vegeta-

tion even during the glacial stages, the moraines being pushed out into

the forests.

In America the Third Interglacial interval is known as the Sanga-

mon; the deposits are composed of dark, black soil which is overlain by

the main or thickest loess deposit of the central United States. There

appears to have been a long interval between the melting of the Third

Glacial ice and the deposition of the loess which contains a terrestrial

molluscan temperate fauna, indicating climatic conditions not greatly

different from those now existing in the same regions (Shimek, 1909).

The geologic deposits of this stage are mainly of three kinds: first,

fluviatile sands and gravels; second, loess; third, hearth or kitchen-mid-

den deposits made by man toward the cold closing period of this stage.

CLIMATIC CHANGES DURING THE THIRD INTERGLACIAL STAGE

The Third Interglacial Stage opens with a renewal or continuation of

climatic conditions favorable to an Asiatic-African fauna exactly similar

to that of the warm Second Interglacial period. This warm fauna is

known as the “Second /. antiquus fauna.” It includes the hippopota-

mus, the straight-tusked elephant (H#. antiquus) and the southern ele-

phant (4. trogontherit). This last species is even referred to by many

writers as #. meridionalis.

OSBORN, REVIEW OF THE PLEISTOCENE 985

This is the Hlephas antiquus Stage of Pohlig, this animal being very

abundant until toward the close of the Third Interglacial Stage when it

makes its last appearance in Europe. The broad-nosed rhinoceros (D.

merckit) is also abundant and appears in Europe for the last time.

The successive climatic phases of mammalian life are most clearly

recorded in connection with the culture stages of the Lower Paleolithic

period, including the pre-Chellean, Chellean, Acheulean, and the begin-

ning of the Mousterian cultures. As indicated on p. 233, the warm

Asiatic-African fauna prevails from the pre-Chellean until toward the

close of Acheulean times, when there is evidence of the advent of a cold

dry continental climate, on the approach of which the hippopotamus,

Elephas antiquus, and Dicerorhinus merckii gradually retreat. Thus at

Villejuif, south of Paris, the late Acheulean implements are found im-

bedded in great drifts of loess, a proof that a cooler, drier climate which

marks the transition from the last warm Interglacial Stage to the Fourth

Glacial advance was prevalent. Chiefly in the southern parts of France

we find the Hlephas antiquus fauna still persisting until the close of the

Third Interglacial Stage or during the early Mousterian period, a sign

that this old African-Asiatic stock did not become extinct but migrated

from central Europe to warmer regions in the south and southwest.

Flora.—tIndications of changes of climate in the Third Interglacial

interval are preserved in the tuf de la Celle-sows Moret (Seine-et-

Marne) ** which overlies Pleistocene river gravels near Paris. The lower

levels contain the sycamore maple (Acer pseudoplatanus), willows

(Saliz), the Austrian pine (Pinus laricia). Higher up in the same de-

posits we find the box tree (Burus) and not uncommonly the fig

(Ficus) ; the canary laurel (Laurus nobilis) occurs less frequently; the

canary laurel and the fig indicate that the winters were mild because

both these plants flower during the winter season. The climate was

more damp and somewhat milder than that of the present time in this

region. The Mollusca of the tufa of La Celle also indicate that the cli-

mate of northern France was more equable so as to permit species now

widely separated to live together. The plants in the highest levels of

the tufa, however, indicate a cooler climate and yield Acheulean flints.

The tufa is itself covered by a sheet of loess corresponding to the return

of a cool, arid period in late Acheulean times.

In Lorraine below the level of the fauna of the Fourth glaciation there

occurs a flora in which the most northerly varieties of the larch (Larix)

and the mountain pine (Pinus lambertiana) predominate. The lignites

of Diirnten and of Utznach near Ziirich (Fig. 9, 18) contain fossil re-

7% HAUG, op. cit., 1908-11, p. 1812.

286 ANNALS NEW YORK ACADEMY OF SCIENCES

mains of forests of Third Interglacial age similar to those which still

flourish in the same region, consisting of spruce, fir, mountain pine,

larch, birches, yews and sycamores with undergrowth of hazel. These

lignitic deposits rest upon the remains of a retreating glacier and are in

turn covered with those of another glacier and are therefore iniergla-

cial.**° With this hardy flora are associated remains of Elephas antiquus,

D. mercku, the urus and the stag.

Fauna.—The mammalian fauna is broadly divided inio: first, the

warm African-Asiatic, which disappears from Europe at the close of the

Third Interglacial Stage; second, the Eurasiatic Forest Fauna, in which

we now include the urus (B. primigenius) and the bison (B. priscus) ;

third. the Tundra Fauna, which retreats afier the Third Glacial Stage

to reappear with the approach of the Fourth Glacial Stage, when the

full tide of Tundra life, including fifteen species of mammals and birds,

and the advance wave of Steppe life, including nine species of mammals

and birds, first arrive in Europe. The chief localities in which the fauna

is recorded are the following:

Warm Stage. CHeLtes™ St. ACHEUL, valley of the Somme, northern France.

Warm Fauna.

Grays THuRBROcK and Inrorp, Essex, England. Warm Fauna.

Cool Stage. TAUBACH-WEIMAR-EHRINGSDORF-ACHENHEIM, Germany. Acheu-

lean Stage. Temperate Fauna.

Dt2gnTeEN, UTzNAcH, near Ziirich. Switzerland (cool flora,

fauna).

LAVISTE. AYGELADES, travertines, Marseilles. France (flora).

Kraprna (cavern of), Croatia (fauna and human remains).

Cold Stage. RrxporFr, near Berlin. Cold fauna.

PILTDOWN MAN, EOANTHROPUS DAWSONI

Fragments of a skull and jaws discovered by Dawson in 1911 near

Piltdown, Sussex, have been described by Dawson and Smith Wood-

ward.*? They were associated in a fluviatile sand layer with a single

pre-Chellean flint and remains of deer (? deer), rhinoceros (D. etruscus

or D. merckit), beaver (Castor fiber), and hippopotamus. The geologic

age is not positively determined by the fauna nor by the nature of the

river gravel deposits in which these specimens were found. The associa-

*® DawkEINs. W. B.: “Classification of the Tertiary Period by Means of the Mam-

malia.” Quart. Jour. Geol. Soc. Vol xxxi, pp. 379-405. Aug., 1880.

® The Chellean culture is placed by Penck and Geikie in the Second Interglacial Stage.

& Dawson, CH., SMITH-WoopWakD. A.. SMITH, G. ELLIoT: “On the Discovery of a

Palzolithic Human Skull and Mandible in a Flint-bearing Gravel Overlying the Wealden

(Hastings Beds) at Pilitdown, Fletching (Sussex). With an Appendix by Prof. Grafton

Elliot Smith.” Quar. Jour. Geol. Soc.. Vol. 69. pp. 117-151. Pis. XV-XXI_ London, 1913.

OSBORN, REVIEW OF THE PLEISTOCENE 287

tion of the pre-Chellean flint and of numerous Chellean flints in the

overlying layer would tend to determine the age as either Third Inter-

glacial or at the earliest Second Interglacial.

The placing of the skull and jaws together as belonging to one indi-

vidual! is not certain, but is highly probable. The cranial bones are

extremely thick; the skull is not Neanderthaloid, but is of a high, dolicho-

cephalic type, with a brain capacity variously estimated at 1,100ccm.

(Elliot Smith, 1913) to 1,500ccm. (Keith, 1913).*** The jaw resembles

closely that of an orang (Simia satyrus); the two lower molar teeth

preserved are more elongated than in any human type; the superior

canine tooth (mistaken by the authors for an inferior canine) resembles

that of the anthropoid ape. Thus the specimen may be concisely de-

scribed as possessing the skull of a man combined with the jaw and the

dentition of one of the higher anthropoid apes. A number of eoliths

and one paleolith were also found in the same layer with the skull.

PRE-CHELLEAN AND CHELLEAN FAUNA

The dawn of the stone industry in Europe is known as the pre-Chel-

lean. It is found at Chelles in France, in Spain and at Piltdown,

Essex, England. It is important to note that the Chellean culture

stage is regarded by Penck, Geikie and others as belonging to Second

Interglacial times, or the Mindel-Riss, while Boule, Haug, Obermaier,

Breuil and Schmidt assign the pre-Chellean-Chellean culture to the

Third Interglacial Stage. While the latter opinion generally prevails

among archeologists there is reason for further investigation before the

geologic age of the pre-Chellean and Chellean cultures can be considered

as definitely determined.

The faunal period of the Chellean culture proper is shown in the val-

leys of the Somme and of the Marne where mingled with the Pre-Chel-

- lean and Chellean flints are found the hippopotamus, the southern ele-

phant (FZ. meridionalis trogontherw) with the straight-tusked elephant

(H#. antiquus) and the broad-nosed rhinoceros (D. merckvi). The typi-

cal site of the Chellean culture stage is near the town of Chelles in the

wide expanse of the Marne valley. The river deposits of the period of

the Chellean culture in this valley are eight meters in thickness and

contain beside the animals named above the giant beaver (Trogonthe-

rium), species of bear, of hyw#na, various kinds of deer, larger and

smaller kinds of wild cattle and a primitive wild horse.

At Abbeville, at the mouth of the Somme, are found remains of the

8iq The corrected and final determination is at 1,300ccm., Smith-Woodward, McGregor.

288 ANNALS NEW YORK ACADEMY OF SCIENCES

same animals associated with those identified as belonging to the sabre-

tooth tiger (Macherodus).

Chellean Culture on the “Low Terraces.”—-\t appears that the Chel-

lean culture stage in many regions was subsequent to the formation of

the terraces; thus Chellean flints may occur in the superficial gravels

both on the “middle” and on the “low terraces.” Haug (1912) sup-

ports the theory that the Chellean culture belongs to the Third Inter-

glacial Stage. ‘This accords with the terrace chronology. The minute

researches of Laville in the basin of Paris are confirmed by the observa-

tions of Commont in the valley of the Somme. Along the Somme Chel-

lean flints occur in the deep gravels overlying the middle and upper ter-

races. ‘The fauna found in the “low terraces” of Chelles and of Grenelle

is the same, namely, H. trogontheru, Trogonthervum cumeri, Hyena

crocuta, D. merchu, Hippopotamus major, Hlephas antiquus. All these

animals found at Chelles occur in the gravels a few meters above the

level of the Marne; they belong exclusively to the sands and gravels at

the base of the diggings in the “‘low terrace.” Similarly in the valley

of the Somme near Abbeville from the base of the “low terrace” are re-

corded Acheulean flints with H. major, D. mercku, Hlephas antiquus,

ete. lLaville collected at Arceuil in the valley of the Biéve, in gravels

assigned to the “low terrace,” Chellean, Acheulean and Mousterian

flints; these “low terraces” are only 5m. above the river level and are

still occasionally flooded with the high waters of the Seine. It is hardly

probable that the close geologic and faunistic association of the Chellean-

Acheulean cultures in these “low terraces” could have been separated by

a very long geologic period, amounting to a hundred thousand years, as

demanded by the theory of Penck.

What is regarded as the typical Third Interglacial fauna of the more

northern regions of Europe as found at Grays Thurrock, Ilford (Hssex,

England) and at T’aubach is as follows:

Grays Thurrock, Ilford (Hssex,

eee eee CauG cere England).— The hippopotamus is

Southern mammoth, H. trogontherii here recorded by Dawkins. The ele-

Broad-nosed rhinoceros, D. merckii phant of Essex is referred by Pohlig

Hippopotamus, H. major to H. trogontheru. The horse

Wild horse, “H. /cabailus (probably © (Hivart) is @onsidered tosnekom=mms

pie er ese Maney the Forest or Nordic type. This

Wild boar, Sus scrofa ferus ‘ ate

: se) n

Been Hacon eres Essex fauna is characteristic of the

Une eos pe eeaae river shores and of the neighboring

Red deer, Cervus elaphus forests and meadows. The lions,

Roe deer, O. capreolus hyenas and bears which are re-

OSBORN, REVIEW OF THE PLEISTOCENE 289

Giant deer, Megaceros corded here are not true cave types,

Hyena, H. crocuta spelwa but are in part ancestors of the cave

Brown bear, Ursus arctos : :

eeiidienat Capra types which appear in the succeed-

Lion, Felis leo antiqua ing reindeer or cavern period.

Wolf, Canis suessi

Badger, Meles taxrus

Marten, Mustela martes

Otter, Lutra vulgaris

Beaver, Castor fiber

Hamster, Cricetus vulgaris

The water vole, Arvicola amphibius

ACHEULEAN CULTURE FAUNA

Warm Stage—The early Acheulean culture as found at Taubach,

Weimar, Ehringsdorf and Achenheim contains the hippopotamus and the

straight-tusked elephant (H. antiquus). The principal feature of the

early Acheulean culture stage seems to be the abundance of these African-

Asiatic animals so that this is commonly known as the “warm Acheulean”

fauna. |

Cool Stage-——The Acheulean culture endured for a long period of time

and toward its close two typical members of the warm fauna, namely,

the hippopotamus and #. trogontherii, disappear. Thus the late Acheu-

lean fauna does not include either hippopotamus or FL. trogontheri but

there still survive the H. antiquus and the broad-nosed rhinoceros (D.

merckit). These animals persisted in Europe for a considerable time

and becoming adapted to a colder climate are sometimes found in asso-

ciation with the advent of the true mammoth (ZF. primigenius) and the

woolly rhinoceros (D. antiquitatis).

During late Acheulean times a dry, cool continental climate prevailed

(Hilzheimer, 1913, p. 145) similar to that of the steppes of southern

Russia between the Ural Mountains and the Caspian Sea. Evidences of

this are observed even in the sheltered valley of the Vézére, a tributary

of the Dordogne in southwestern France. Similarly as regards north-

central France, Obermaier (1912, pp. 122-124) observes that while the

climate was mild and temperate and the country still forested at the be-

ginning of the Acheulean culture, in late Acheulean times the implements

at Villejeuf south of Paris, are found embedded in drifts of loess, a proof

that the colder climate which marks the transition from the Third Inter-

glacial Stage to the Fourth Glacial Stage was now beginning to prevail.

The fauna is still that of FH. antiquus and D. merckii. More rarely

(Schmidt, 1912) Acheulean paleoliths are associated even with remains

990 ANNALS NEW YORK ACADEMY OF SCIENCES

of the woolly mammoth and the woolly rhinoceros, indicating that in

northern localities the Acheulean culture reached the cold period of the

Fourth Glacial Stage.

Krapina Neanderthaloid Race-——-To the Acheulean Stage there is re-

ferred a human tooth found at Taubach. Of much greater importance

is the presence of abundant skeletal remains of men of a primitive Nean-

derthaloid race found in the cave-shelter of Krapina in Croatia. These

remains are positively associated with the Acheulean stage by Schmidt

(1912, p. 256) but they are regarded as more recent, of the late Mous-

terian culture stage; by Breuil. The remains as finally described by

Gorjanovic-Kramberger®? include hundreds of human bones intermingled

in various separate strata with hundreds of stone implements and chips

and thousands of animal bones. Of the contemporary fauna are re-

corded Ursus speleus, Bos primigenius, Equus ? caballus, Dicerorhinus

mercku, Megaceros euryceros, Castor fiber, Arctomys marmotta. The

human racial type is unquestionably related to that found at Neander-

thal and Spy. The race is somewhat dwarfed, of broader head form,

with less prominent supraorbital processes. The species is Homo nean-

derthalensis.

Mousterian Culture, Temperate Fauna—The earliest strata of the

Mousterian culture stage in France show a fauna not differing essen-

tially from that of the late Acheulean stage, namely, a fauna containing

Elephas antiquus and Dicerorhinus merckii. Thus in La Micoque, one

of the oldest stations in the Vézere valley, Dordogne, in which the eul-

ture belongs to the transition between late Acheulean and early Mous-

terian times, in the very lowest layers are found traces of the broad-

nosed rhinoceros (D. merckiw) associated with remains of the moose

(Alces). But the last glacial stage is approaching and D. merckw gives

place to the migrants from the tundra region of the northeast, covered

with hair, adapted to an arctic climate, namely, the mammoth and the

woolly rhinoceros. The main succeeding portion of the Mousterian cul-

ture was contemporaneous with the Fourth Glacial Stage and the cold

tundra, steppe fauna.

82 GORJANOVIC-KRAMBERGER, KARL: “Der Diluviale Mensch von Krapina in Kroatien

Ein Beitrag zur Paidoanthropologie. Studien iiber die Entwickelungsmechannik des

Primatenskelettes mit besonderer Beriicksichtigung der Anthropologie und Descendenz-

lehre.”” Herausgegeben von Dr. Otto Walkhoff. C. W. Kreidel’s Verlag, 4to. Wiesbaden,

1906.

OSBORN, REVIEW OF THE PLEISTOCENE 29]

FourtH GLACIAL STAGE—WURM, MECKLENBURGIAN, WISCONSIN

BEGINNING OF THE REINDEER AND CAVE PERIOD

The Fourth Glacial Stage, like the First, is believed to have been

nearly contemporaneous in Europe and North America, consequently the

estimates of Postglacial time in one country have an important bearing

on the other. The First Maximum of the Fourth or Warm glaciation in

the Alps is estimated by Penck as occurring 40,000 years ago. It was

followed by the slight recession known as the Laufenschwankung, a tem-

perate retreat followed in turn by the Second Wiirm Maximum, which is

estimated as occurring 20,000 years ago. Similarly in America the

“early Wisconsin” is followed by a recession interval (Peorian), and

this in turn by the “late Wisconsin” which is the final great glaciation in

America. The contemporaneous Mecklenburgian of the North German

lowlands gave rise to the “Upper Drift,’ which in some respects bears a

striking resemblance to the Wisconsin Drift of America both in its sys-

tems of moraines and in its topography. This stage also includes appar-

ently the “Upper Drift” of northern England with which the drift of

the Alps correlates well. The Upper Drift of England covered also a

large part of Wales. In Germany glaciation also occurred in the Rie-

sengebirge and the Black Forest.

In America part of the “Upper Drift” is loess-covered and in the opin-

ion of Koken and Schmidt the Upper Drift of Germany is also partly

covered with the “Newer Loess.” The Postglacial Stage did not exhibit

a steady amelioration of climate after the culmination of the Fourth

Glacial Stage, but there is evidence of great oscillations and renewed

glacial advances both in northern Britain and Scandinavia and in Ger-

many. These Postglacial advances, as most clearly defined in the Alpine

region, have been termed by Penck the Buhl, the Gschnitz and the Daun.

They are correlated by paleontologists and anthropologists quite closely

with the successive faune and archeological implements of Postglacial

time.

Period of the Final Glacial Maximum.—The length of time which has

elapsed since the close of the Fourth great glacial advance is estimated

in America by the recession of the Falls of Niagara. This recession

began with the end of the Wisconsin glaciation which is believed to have

been contemporaneous with the Wtirm. As early as 1829 Bakewell esti-

mated that since the Falls of Niagara were receding by the erosion of the

Niagara gorge at the rate of three feet annually about 10,000 years had

elapsed since the end of the Glacial Epoch. Lyell visited Niagara in

1841 and after consideration of all the data of erosion concluded that

909 ANNALS NEW YORK ACADEMY OF SCIENCES

the time since the last great glacial stage was not less than 31,000 years.

Gilbert, Upham and other geologists of the United States Geological

Survey aiter pointing out many sources of error im all such calculations

were inclined to the adoption of periods ranging from 6,000 io 10,000

years. Gilberts survey is the most careful which has been made; he

estimated (1896) that the gorge of Niagara is not more than 7,000 years

of age. The Canadian geologist Spencer arrived at a result almost iden-

tical with that of Lyell, namely, 32,000 years.

The most careful estimates on the subdivisions of Postglacial time in

Europe are those of Penck (1909, p. 1168) which may be briefly sum-

marized as follows:

IV. 2nd WtUeM wWAXxIMUtM,. beginning of Upper Palzxo-

lithic .culivtes: < 2.45) epee ee eee ee 20.000 to 34,000 years

Achen reireat. period of Aurignacean and Soluirean

CUlGMTE «oooh oak feces So eee eee eee

Biihl advance. period of early Magdalenian culture

( NGHSER) uF oe es daw bonis teed ee eee 16.000 “ 24000 *

Post-Biihl, period of late Magdalenian culture..... 10.000 * 16,000 “*

Daun Stage. period of the close of the Upper Palzo-

lithic, Azilian culture......... tiasip cece ee 7.000 .

Azve. of Capper in: Marope. nF ven en eeeees 4.000 * 5.000 *

According to Heim’s** calculations the period since the deposition of

the Biihl moraines in the Lake of Lucerne amounis approximately to

16,000 years and if the Magdalenian culture deposits of this formation

are of the age of the Buhl advance we my estimaie with Neusch that

the Buhl advance occurred at least 24,000 years ago. This advance is a

very important period because it represents the last of the Arctic, Tundra

and Steppe Faunas in central Europe prior io the establishment of the

modern forested conditions and fauna.

Fauna of the Fourth Glactal Stage-——The severe climate of the Fourth

Glacial Stage is indicated by the mammalian life found at Achenhemm

and Sirgensiein associated with what is known as the “full Mousterian”

Palzolithic culture (Schmidt, 1912). Here for the first time there is a

predominance of the Tundra Fauna (fifteen species) as well as the ad-

vent of the Steppe Fauna (two species), while the Forest Fauna (seven -

‘species) and Meadow Fauna (four species) are still represented. The

straight-tusked elephant (£. antiguus) and broad-nosed rhmoceros (D.

merckit) have disappeared and are now replaced by the woolly elephant

(E. primigenius) and woolly rhinoceros (D. antiquitatis). The mam-—

= Heim, A.: “Ober das absolute Alter der Eiszeit.” Vierteljahrsschrift der natur-

forsch. Ges. in Ziirich, Jahrg. 39. No. 2. pp. 180-186. 1894.

= : _ aaa ™

OSBORN. REVIEW OF THE PLEISTOCENE 293

mal life of the Fourth Glacial Stage at Achenheim and Sirgenstein near

the upper Rhine is divided among the various faune as follows:

TUNDRA

Blephas primigenius After the First Maximum of the

eae Fourth glaciation the Laufen-

Rangifer tarandus

schwa ay 1porarily

Vulpes lagopus chwankung may have temporarily

Lepus variabilis favored the return of the Llephas

Myodes torquatus antiquus and D. merckii fauna as

STEPPE far as northern France because we

Equus germanicus (Steppe type)

occasionally find a glacial mixed

Spermophilus rufescenus

fauna where FH. antiquus and D.

FOREST mA ; = ;

Gees ciaphas merckw occur in association with L.

Lynchus lynr primugenus. The close of the

Canis lupus Fourth Glacial Stage or Second

Vulpes aloper

Arvicola amphibius

Ursus spelevus

Wiirm Maximum is marked by the

first appearance of very numerous

Geis arctic rodents, especially of the

Arctomys marmotta banded lemming (Myodes torqua-

Ibex sp.? tus) type, which constitute the so-

Sede Os called “Lower Rodent” layer. The

EADOW - °

Bison priscus animals ( Schmidt, 1912) character-

Bos primigenius istic of this Lower Rodent, stratum

ASIATIC as found at Sirgenstein, Wild-

Hyena spelea scheuer and Ofnet are as follows:

Felis leo spelea

TUNDRA

This “Lower Rodent?’ stratum Canis lagopus

Fetorius erminea

Lepus variabilis

Myodes obensis

with a fauna such as we find at pres-

ent in far northern Russia registers

the coldest climate of Pleistocene “ — torquatus

times, corresponding to the Second Lagopus albus

Maximum of the Fourth Glacial oRrst

Stage. It is well known that this ee eee

MEADOW

Lower Rodent fauna is not local but fo Eas

rather a widespread phenomenon ex-

tending over northern and southern Germany and Belgium (Schmidt,

1912, p. 261). The presence of this Lower Rodent fauna at Thiede near

Braunschweig in the border region of the Upper Glacial Drift of Germany

is of special significance, as Wahnschaffe observes. This is the classical

locality for lemming, the remains of lemmings being associated solely with

those of arctic fox, arctic hare. reindeer, musk ox and mammoth.

9G4 ANNALS NEW YORK ACADEMY OF SCIENCES

Upper Sands of Mauer—In the Upper Sands of Mauer, Wurm (op.

cit. 1913) and Forster** have observed the presence of a sub-Arctic cold

fauna including the arctic reindeer (Rf. tarandus) and the banded lem-

ming (Myodes torquatus), the steppe suslik (Spermophilus rufescens),

also the steppe weasel (Fetorius eversmanni). This is positively corre-

lated with the fourth glaciation by Wurm.

The fauna of the Wildkirchli cave on Mont Centis in Switzerland

shows no typical interglacial forms and may indicate either the approach

of the Wiirm Glacial period or its Laufenschwankung (Schmidt, 1912,

p. 193)"

Mousterian Paleolithic Culture-—The Mousterian Paleolithic culture

of the Neanderthaloid race appears to have begun toward the close of the

Third Interglacial Stage and continued throughout the entire period of

the Fourth Glacial, including the First Maximum, the Laufenschwan-

kung, and the Second Maximum, ending with the “Lower Rodent” fauna

in the deposits of the “Newer Loess.” Thus Koken and Schmidt speak

of the Mousterian culture as found at the base of the “Younger Loess.”

The early men of the Mousterian culture period witnessed in the north

the Elephas antiquus and broad-nosed rhinoceros, and the hippopotamus

in the south, as proven in the Prince’s Cave of Monaco. The culture

lasted through all the severe climatic conditions of the entire Fourth

Glacial Stage. The men of the Neanderthal race in Mousterian times

sought the rock shelters and grottos or entrances to caverns so that the

greater number of paleoliths are found mingled with the remains of ani-

mals of the chase around the old hearths. The commonest game evi-

dently consisted of the wild horse, urus and reindeer. Less frequently

the ibex and stag were objects of the chase.

Neanderthal Races——To the Neanderthal race (Homo neandertha-

lensis,*°) in addition to the remains of Krapina, belong many and widely

distributed remains, including the classic type of Neanderthal discovered

in 1856, the Gibraltar skull discovered in 1848, those of La Naulette,

Spy (1887), Krapina (1902), Malarnaud, La Quina, Le Moustier

(1908) and La Ferrassie. The skull of Neanderthal man was character-

& FORSTER, F.: “Uber eine diluviale subarktische Steppenfauna aus den Sandhtgeln

von Mauer.” YVerhandl. naturw. Verein. Karlsruhe, 1913.

8 The mid-Pleistocene man was definitely named Homo neanderthalensis by Cope in

1893 at the time of the discovery of the skeletons of Spy; it seems, however, that King

had previously (1864) used the same term. Schwalbe (1901) remarks, . = **the-

species of man which I, together with King (1864) and Cope (1898), dealentined as.

Homo neanderthalensis.’”’ In the following year (1902) the same author introduces the

term Homo primigenius, which is that generally adopted in Germany. Among French

authors the same man is known to-day as Homo mousteriensis. This polynomial usage

serves at least to emphasize the unanimous opinion as to the distinct specific character

of mid-Pleistocene man.

OSBORN, REVIEW OF THE PLEISTOCENE 295

ized by an extremely receding forehead, by the great prominence of the

supraorhital ridges. and by a rather slender jaw. The occipital projec-

tion for the attachment of the superior muscles of the neck was large,

indicating that these muscles at the back of the neck were strongly devel-

oped, a character necessary to meat-eaters before the invention of knives

Fic. 16.—Neanderthal skull

Skull of a Neanderthal man from the cavern of La Chapelle-aux-Saints (Corréze), France.

After Boule.

and forks. This primitive type of man was shorter than the average

European (that is, 5 feet 814 inches) ; he is estimated of as low stature

as 5 feet 314 inches. His lower limbs were especially powerful, but his

gait seems not to have been fully erect, for the knees are bent slightly

forward.

296 ANNALS NEW YORK ACADEMY OF SCIENCES

The most remarkable skull and skeleton of Mousterian age is that

(Fig. 16) found by the Abbés J. and A. Bouyssonie and L. Bardon in

the cavern of La Chapelle-aux-Saints (Corréze) in 1908, associated with

stone implements and remains of the reindeer, urus, ibex and woolly

rhinoceros. The cranium is dolichocephalic, with prominent supra-

Se, ve CR OKS GAT

Fic. 17.—Neanderthal man

Reconstruction of the head of Homo neanderthalensis by Charles R. lxnight under the

direction of the author. 1910.

orbital processes and relatively short and broad nose, weak lower jaw,

lacking the prominent chin process. These characters, as well as the

posterior position of the foramen magnum and the form of the palate,

are distinctly simian or pro-human.*‘®

POSTGLACIAL STAGE—CONTINUATION OF UPPER PALXOLITHIC,

REINDEER OR CAVE PERIOD

CLIMATE

For a long period the fauna of Postglacial time in Europe remained

practically the same, namely, during the Mousterian, Aurignacian, Solu-

trean and Magdalenian culture periods. The cold fauna is shown both

in the animal remains and in the art, which is so characteristic of

Aurignacian and Magdalenian times.

As the “Lower Rodent” layer of Mousterian times is referred to the

Second Maximum of the Fourth glaciation and the period of most intense

3 BouLE, M.: “L’'Homme Fossile de la Chapelle-aux-Saints.”” L’Anthropol.. Vol. XIX,

pp. 519-525. 1909.

OSBORN, REVIEW OF THE PLEISTOCENE 297

cold it is followed by an amelioration of climate (Achenschwankung)

during Aurignacian and Solutrean times, which in turn is succeeded by

a recurrence of a colder climate during the Bihl advance, at which stage

it is believed the “Upper Rodent” layer of Magdalenian times indicates

a fresh invasion of Arctic-Tundra types. This tundra “Upper Rodent”

layer occurs in the deposits of Sirgenstein, Wildsheuer and Ofnet; it is

associated along the upper valley of the Danube with early Magdalenian

paleoliths, and continues in Hohlefels, Schussenquelle, Andernach, Miun-

zingen and Wildscheuer in association with the Middle Magdalenian cul-

ture stage; there is, moreover, in Hohlefels, Schmiechenfels and Propst-

fels an association of the cold steppe fauna with a late Magdalenian cul-

ture stage. After this there was a decided amelioration of climate, for

in the final Azilian-Tardenoisian stage all the Steppe, Tundra and Alpine

types disappear.

UPPER PALZOLITHIC, FOUR OR FIVE HUMAN RACES

(1) The Grimaldi Race, a negroid type is found close to the warm

faunal level at Mentone. (2) An initial feature of the long Postglacial

Stage is the entire disappearance of the Neanderthal race of men and

the invasion of Europe by a new race of the much higher Crd-Magnon

type referred to the existing human species (Homo sapiens). This

race was discovered in Dordogne at the hamlet of Cro-Magnon from

which it takes its name. It has also been found in the caverns near

Mentone lying above remains of the Grimaldi race. Cré-Magnon re-

mains are also found at several localities in France in deposits of Aurig-

nacian and Magdalenian times. In the south in Aurignacian times this

was a tall, well-proportioned, hunting and nomadic race; in the more

severe climate of the north it occurs as a dwarfed race. The skulls are

dolichocephalic, with very broad cheek bones, hence designated as dishar-

monic. The brain capacity is equal to that of the early Neolithic races

and surpasses that of many modern races. It is probable but not posi-

tively demonstrated that to this Cré-Magnon race we owe the early arts

of engraving, sculpture, design and mural painting which are preserved

in the Aurignacian and Magdalenian deposits of France and northern

Spain. (3) In Solutrean times there was at Briinn and Prédmost (Mo-

ravia) another dolichocephalic race which is considered as distinct or

transitional because unlike the typical Cré-Magnon race the cheek bones

are narrow and the skull is thus harmonic. This is the Briinn-Prédmost

race. (4) After the close of the Magdalenian culture stage, or during

the so-called Azilian-Tardenoisian culture, the first brachycephalic hu-

man races make their appearance in Europe. Their remains have been

998 ANNALS NEW YORK ACADEMY OF SCIENCES

found at Ofnet, Furfooz and at Grenelle. This is commonly known as

the Grenelle or Furfooz race; it is very distinct from the preceding races

in bodily structure and in culture.

Aurignacian, First Upper Paleolithic Culture Stage—This first of

the Upper Paleolithic culture stages is widely distributed in western

Europe. It takes its name from the small grotto of Aurignae (Haute

Garonne) where the first discoveries of the culture and of a number of

skeletons were made in 1852. The arts of engraving on bone and stone,

of drawing and painting in single lines, of sculpture of the human and

of animal figures, all in bold but primitive forms, first appear in Aurig-

nacian times. Thus man through his art begins to make a permanent

record of the contemporary mammalian life, especially of the mammoth,

bison, reindeer and cave bear. With early Aurignacian times the cold

climax is passed but we still find remains of the Arctic lemming (Myodes

torquatus) fauna. The mammalian list of the Aurignacian stations both

of the “Newer Loess” and of the caverns still gives a cold aspect with its

Tundra-Steppe-Alpine types with which no warmer types are associated.

In middle and late Aurignacian times the lemmings for a time disappear ;

otherwise the fauna retains its northern character (Gulo luscus, Lagopus

alpinus), which is not essentially altered by the presence of the hyena

and stag.

Solutrean, Second Culture Stage-——This stage, which takes its name

from the type station of Solutré (Sadne-et-Loire) represents the climax

of perfection in the Upper Paleolithic flmt mdustry, which appears to

represent partly a development of Aurignacian workmanship and partly

a culture invasion. With Solutrean times Schmidt correlates the three

Briinn skeletons and Prédmost (Moravia). It is noteworthy that no evi-

dences of a Solutrean art have been discovered. The fauna like that of

the Aurignacian represents an amelioration of the extreme cold of the

Fourth Glacial maximum. The wild horse and reindeer are abundant

as well as the mammoth, rhinoceros, wolf and cave bear. Perhaps the

somewhat more frequent appearance of such cold faunal types as the

Alpine hare and grouse betoken the approach of the colder Buhl stage

of Magdalenian times.

Magdalenian, Third Culture Stage and Fauna—This third Upper

Paleolithic culture takes its name from the station of La Madeleine

(Dordogne). It is distinguished by decline in the perfection of the flint

industry as compared with the Solutrean stage, by a very decided devel-

opment of bone implements, and by a surprising advance in the arts of

engraving and painting and the sculpture of animal forms in bone and

ivory. The Magdalenian stage corresponds with the “Upper Rodent”

OSBORN, REVIEW OF THE PLEISTOCENE 299

strata which registers a period of extreme cold correlated with the Bihl

glaciation (Penck, Schmidt), which again attracts the cold-loving ro-

dents. The “Upper Rodent” strata associated with Magdalenian cultures

in the shelters of Schweizersbild and Kesslerloch are of more recent date

than the neighboring “Lower Terraces.” This “Upper Rodent” stratum

contains an abundance of Tundra and Steppe types of smaller rodents

and represents the last stage of extreme cold in Europe. Thus the

banded lemming (Myodes torquatus) corresponds with early Magda-

lenian times. In the upper levels of the Upper Rodent layer, which be-

long to late Magdalenian times, the Tundra fauna gradually gives way

to a more abundant Steppe fauna, the banded lemmings becoming less

frequent while the jerboas (Alactaga jaculus), the hamsters (Cricetus

pheus), the susliks (Spermophilus rufescens) become more abundant.

The reindeer and the wild horse are very abundant. The mammoth, the

woolly rhinoceros and the cave bear gradually retire from the middle and

southern mountains of Germany, and in the very highest Magdalenian

culture layers the fauna begins to approach that of recent times, namely,

the Eurasiatic Forest fauna. In Schmidt’s opinion there is no question

as to the similarity of age of the Magdalenian layers of the Miinzingen

loess deposits with the cave deposits of Schweizersbild and Thaingen.

From this evidence it can be positively determined that the chief deposits

of the “Newer Loess” occurred after the Fourth glaciation.

With Magdalenian times are associated the skeletons of La Madeleine,

Laugerie Basse, Chancelade, La Hoteaux and Duruthy; all are regarded

as of the Cré-Magnon racial type.

POSTGLACIAL FAUNA

The mammal fauna of this long Postglacial period is the same in the

upper valleys of the Rhine, the Danube, the Dordogne and the Garonne.

It extends throughout the Pyrenees and the Cantabrian Alps of northern

Spain. Even the reindeer invaded this region (Harlé) but the stag (C.

elaphus) is more abundant. The moose (Alces) also invaded the Pyre-

nees and northern Spain. The Saiga antelope (S. tartarica) occurs at

thirteen localities in southern France (Harlé), and the steppe suslik

(S. rufescens) is very abundant.

It is a grand assemblage of the European Forest and Meadow types

mingled with a few Eurasiatic-Alpine types, abundant Eurasiatic Forest

types, but with the Tundra and Steppe types predominating numerically

until the close of the Magdalenian period, when the Forest types again

begin to greatly predominate. The numerical succession in Germany and

Austria may be tabulated from the invaluable tables presented by

200 ANNALS NEW YORK ACADEMY OF SCIENCES

Schmidt. We owe our knowledge of the cold fauna of southern France

chiefly to Harlé (1871-1912).

wn ee. mM n 2

ae = a ey =

= 0 oo oo o°o +o

Se 22) £2) 33s

= nD Qa 2a San a

: es WE se Lene wh < = =

Postglacial, Daun Stage, Azilian-Tardenoisian Cul-

ture: ‘OTmet, Tstei 22-2 fa. Ase eee ee Gee i

Postglacial, Gschnitz Stage, Hohlefels, Late Mag-

dalenian Cmiaeeycie. fos a oe oe ee eee eae ae T 6 i 5 i2

Fostglacial, Middle Magdalenian Culture, Upper

Rodent: Siratass. one k «see eee cree 10 7 it 6 S

Postglacial, Early Magdalenian Culture, Biibl

Stace; Sinsensteimnhssd Lo ta eit Moers ae ae 9 5 5 ie 3) 3

Postglacial, High Magdalenian Culture, Thaingen. 15 14 ok 13 21

Postglacial, High Magdalenian Culture, Schwei-

WETSUIT: one oe Chip we wae Gale eee ee er ee ee T = ee 6 8

Postglacial, “Solutrean. Ciltavels 3-25. eee oe eee 6 3 a ole

Postglacial, Late Aurignacian Culture........... 10 4 ot: 1 >

Postglacial, Aurignacian Culture, Sirgenstein.

Omet lai wale Cem ae fe des eee Oe eee 7 cb 1 ab 3

IV. Glacial Maximum, Lower Rodent Layer..... i 4 z 3

VY. Glacial Stage, Mousterian Culture........... Ss 5 1 Z 4

The fauna of Voklinshofen includes a similar intermingling of tundra,

steppe, mountain, and meadow-forest types. The same is true of the

scattered deposits’ in Thuringia near Saalfeld, Gera, Jena, Leipzig, ete.

The loess fauna near Wurzburg, Bavaria,** also includes twenty species

of mammals divided into typically modern tundra forms of northern

Asia, typical modern steppe forms of central Asia and Siberia, together

with the four characteristic great mammals of the period, the mammoth,

the woolly rhinoceros, urus and bison. The arctic character of the fauna

-of Chateauneuf-sur-Charente*® in central France is very conspicuous,

most of the species belonging either to the tundras or the steppes of

modern Europe. The bones of many young animals occur in this deposit,

which may be explained perhaps on the supposition that the animals fell

into the fissure while the opening was lhghtly covered with snow, the

young being the most frequently entrapped. Among the chief localities

where this grand Fourth Glacial and Postglacial fauna have been dis-

covered are the following:

PoHuic, H.: ‘“‘Vorliufige Mittheilungen tiber das Plistocen, insbesondere Thiirin-

gens,’’ Sitzungsber. Niederrhein. Ges. Bonn, pp. 2-15. Mar. 3, 1884.

88 NEHRING, A.: “‘Ubersicht tiber vierundzwanzig iain eas Quartar-Faunen,”’

Zeitschr. deutsch. geol. Ges., pp. 468-509. Jahrg. 1880.

8° BouLE, M., and CHAUVET, G.: “Sur l’existence d'une faune d’animaux arctiques

dans la Charente a4 l’époque quaternaire,’ C. R. Acad. Sci. Paris, Vol. XXVIII, pp. 1188-

1190. 1899.

OSBORN, REVIEW OF THE PLEISTOCENE 301

Localities Culture Zones

KESSLERLOCH Cave, near Thaingen (Fig. 9, 37)

SCHWEIZERSBILD Cave, near Schaffhausen (37)

Scattered deposits in Thuringia, in northern Germany,

Saalfeld, Gera, Jena, Leipzig (29, 30, 31)

Wwtrzpure, Bavaria, Loess deposits (33)

SWABIA and FRANCONIA, cave deposits

VOKLINSHOFEN, Alsace (28)

Magdalenian Stage

Aurignacian and Solu-

trean Stages

Solutrean Stage

Aurignacian and Solu-

trean Stages

MONTMAURIN Cave (Haute-Garonne), Upper levels (35) Magdalenian Stage

CHATEAU NEUF-SUR-CHARENTE (Charente) (36)

The large mammals of the period are more fully known than in any

previous Pleistocene stage both through paleontological researches, which

date back to the beginnings of the science of vertebrate paleontology in

Germany and France, and through the extraordinarily accurate carvings

and drawings in the caverns of Dordogne and northern Spain. These

drawings have been reproduced with remarkable fidelity, chiefly by

Breuil.°° The chief elements in the larger mammal fauna were as fol-

lows:

FOREST AND MEADOW FAUNA

Schwerzershild Cave (Fig. 9,

Moose, Alces palmatus

Zeitschr. Deutsch. Geol. Ges., pp. 468-509.

: “Die kleineren Wirbeltiere vom Schweizersbild bei Schaffhausen,”

allg. schweiz. Ges. gesiim. Naturwiss., Vol. xxxv, pp. 41-77.

Persian deer, Cervus maral

Red deer, Capreolus capreolus

Roedeer, Cervus elaphus

Wild cattle, Bos primigenius

Bison, Bison priscus

Forest horse, Hquus caballus

Bear, Ursus speleus

Lion, Felis leo spelea

Brown bear, Ursus arctos

Wild boar, Sus scrofa

TUNDRA FAUNA

Woolly rhinoceros, Diceros antiqui-

tatis

Woolly elephant, Elephas primigenius

Musk ox, Ovibos moschatus

Reindeer, Rangifer tarandus

Arctic fox, Vulpes lagopus

Elasmothere, Elasmotherium sibiri-

cum

37).—The Magdalenian, or rein-

deer man apparently arrived in

the Schaffhausen region long

after the maximum Fourth gla-

ciation, during the Biih] advance,

the period of deposition of the

“Upper Rodent” layer with its

cold Arctic and Steppe fauna.

The deposits of the Schweizers-

bild cave as recorded by Neh-

ring®*? present the fullest succes-

sion and extend over a very long

period of time as exhibited in the

following layers:

Neolithic :

5. Gray culture layer, forest fauna.

9 See publications of Abbé Ilenri Breuil, Marcellin Boule, Le Comte Begouen, I’.

Bourrinet, L. Capitan, Emile Cartailhac, Lalanne, Lartet and Christy, Déchelette, Hugo

Obermaier, Péyrony, Reinach. Others are in preparation.

% NHHRING, A.: “itbersicht tiber vierundzwanzig mitteleuropiische Quartir-Faunen.”

1880.

Denkschr.

1896.

)

305 ANNALS NEW YORK ACADEMY OF SCIENCES

STEPPE FAUNA Paleolithic, Magdalenian :

Saiga antelope, Anielope saiga 4. Upper Breccia, or “Upper Ro-

Asiatic wild ass, Hquus hemionus dent” layer, steppe fauna.

Wild horse, EB. przewalskii ? sp. 5. Yellow culture layer, palzolithic

Jerboa, Alactaga jaculus “Reindeer Age,” steppe fauna.

. Lower Breccia. or “Lower Ro-

dent” layer, animal remains

ALPINE FAUNA and traces of man, tundra

Ibex, [bez priscus fauna.

Chamois, Rupicapra 1. Diluvial layer. No fossils.

Desert horse, E. caballus celticus

(2) Of these the “Lower Rodent” layer contains a pure arctic tundra

fauna, such as the vole, hare, fox, the reindeer, the ptarmigan. There

are no traces of man. In the layer above these the early steppe animals

begin to appear, the hamsters and picas. (3) Then in the “Yellow Cul-

ture” layer there is an assemblage of pure steppe forms, susliks, dwarf

picas and wild horses, all pointing to the absence of forests; but at the

top of this layer the first squirrel (Sciwrus) appears as the harbinger of

forests. (4) In the “Upper Rodent” layer the steppe fauna begins to be

intermingled with an increasing number of forest types, such as squir-

rels, dormice, and the pine marten. (5) Finally we reach the “Gray

Culture” layer, composed of the modern forest dwellers, such as the squir-

rel, the beaver, the pine marten, the stag, the roe, the wild boar, the

brown bear. The “Lower Rodent” layer is contemporaneous with the

Mousterian culture, while the “Upper Rodent” layer belongs to Magda-

lenian times. The uppermost “Gray Culture” layer with its forest fauna

belongs either to the closing Paleolithic or to Neolithic times.

Kesslerloch Cave.—Similar conclusions result from the study of the

geologic conditions surrounding the Kesslerloch Cave of Thaingen in

Switzerland (Fig. 9, 37). This famous cave lies on the edge of a

moderately wide valley, traversed by a brook.** In this sheltered, well-

watered, hilly region, woods flourished and harbored the forest animals,

at the same time that the glaciers retreating southward left damp and

stony areas, closely followed by a tundra fauna. The woolly rhinoceros —

and the mammoth persisted longer here than in other parts of Europe.

As analyzed by Niiesch, we discover here mammals distinctive of the tun-

dras, of the steppes, of the modern Alps (marmot, chamois, ibex), of the

meadow-forests (bison, urus), and finally of the modern forest type (lion,

wolf, brown bear, pine marten, squirrel, wild boar, and stag). These

mammal zones undoubtedly correspond with the passing or evolution of

82 NUESCH, J.: “‘Das Kesslerloch, eine Hoéhle aus paldolithischer Zeit. Neue Grabungen

und Funde.’”’ Neue Denkschr. allg. schweiz. Ges. gesam. Naturwiss., Vol. XXXIX, Pt. 2,

pp. 1-72. 1904.

OSBORN, REVIEW OF THE PLEISTOCENE 303

several human culture stages (perhaps the Aurignacian, Solutrian, and

Magdalenian). While the tundra fauna was pushing southward into

the heart of Switzerland, it had already vanished from central Germany,

Belgium and France, where it had been superseded by a. steppe, or even

a meadow-forest fauna. ‘The human artifacts show that these deposits

are contemporaneous with those of Schweizersbild, both belonging to

Magdalenian times. A hearth, with ashes and coals, and many charred

bones of old and young mammals, including the woolly rhinoceros, have

been found here. The human remains show that a race of pigmies dwelt

here smaller even than the small men of Schweizersbild, their height

being estimated at 120cm. (4 feet).°* The horse of Kesslerloch shows

many resemblances to the Przewalsky horse of the high steppes of Cen-

tral Asia.®*

It is characteristic of these faunas that among species still living are

mingled remains of the great extinct mammals of the times. Another

feature is that occasionally the Steppe, Tundra and Forest faunas are

found either nearly pure or entirely distinct and separate as in the lower

deposits of Thiede near Braunschweig, above cited. More often as in

Schweizersbild and Kesslerloch they are successive or superposed upon

each other. |

Beside the cavern deposits are those in the loess. Thus in the “Upper

Loess” near Wiirzburg, Bavaria, Nehring®® has recorded both a Tundra

and a Steppe fauna, including beside the still living types the woolly

rhinoceros, the mammoth, the urus and the bison.

MIGRATIONS OF THE LARGE MAMMALS OF THE FOURTH GLACIAL AND

POSTGLACIAL PERIOD

Over the greater part of the Iberian Peninsula the stag (Cervus

elaphus) took the place of the reindeer. There is no trace of the en-

trance of the Steppe Fauna at any period into Spain or Portugal. The

Pyrenees also presented a barrier to the greater part of the tundra

fauna, yet the Norwegian lemming (Myodes lemmus) penetrated Por-

tugal to the vicinity of Lisbon. The cold fauna (FZ. primigenius, E.

antiquitatis, U. speleus, F'. spelea) is not represented in Portugal, but

EH. primigenius has been discovered in two localities on the extreme

northern coast of Spain, in the Province of Santander bordering the

Bay of Biscay. D. antiquitatis also occurs in the same _ province.

% NUESCH : op. cit., p. 21.

%4 StupeErR, T.: “Die Knochenreste aus der Héhle zum Kesslerloch bei Thayngen,” Neue

Denkschr. allg. schweiz. Ges. gesam. Naturwiss., Vol. XXXIX, Pt. 2, pp. 73-112. 1904.

% NEHRING, A.: “itbersicht tiber vierundzwanzig mitteleuropiiische Quartir-Faunen,”

Zeitschr. Deutsch. Geol. Ges., pp. 468-509. 1880.

304 ANNALS NEW YORK ACADEMY OF SCIENCES

Rangifer tarandus is found im the cavern of Serinya south of the Pyre-

nees (Torralba). It also has been recorded recently (Obermaier) in

the cavern of Altamira, near Santander. The Alpine chamois (Rupi-

capra) occurs south of the Pyrenees and the ibex is traced to Gibraltar.

Thus Harlé*® concludes it is certain that the “cold fauna” of France

spread along the northern coast of Spain flanking the Pyrenees into

Catalonia, including the mammoth, reindeer, chamois, woolly rhinoceros,

and spreading as far west as Santander. This is also the range of the

Cr6-Magnon race of men.

Mammoth.—The woolly mammoth (££. primigenius) now reaches the

height of its evolution and specialization. As preserved in the frozen

tundras of northern Siberia and as represented in very numerous draw-

ings and engravings by the Upper Palzolithic artists, it is the most com-

pletely known of all fossil mammalia. Its proportions as shown in the

accompanying figure, which represents the information gathered from

all sources, are entirely different from those of either the Indian or

African elephant. The head is very high and surmounted by a great

mass of hair and wool. Behind this is a sharp depression separating the

back of the head from the great dorsal hump. The hinder portion of

the back falls away very rapidly and the tail is short. The overcoat of

long hair nearly reaches the ground, and beneath this is a warm under-

coating of wool.

As described by Salensky** from the wonderfully complete specimen

discovered in 1901 on the banks of the Beresowka River in northeastern

Siberia. this animal developed characters which absolutely exclude the

possibility of its ancestry or relationship to the existing Indian elephants.

The hind foot was four-toed, or tetradactyl, and not five-toed as in the

living forms. The head was larger as compared with the length of the

body than in recent elephants, a character which stands in close connec-

tion with the enormous development of the tusks; these were distin-

guished by their spiral form, the points directed inward. The ears were

very small and covered with hair. The tail was relatively shorter than

in the existing elephants and was provided with a tassel of long, bristly

hair at the end. The color of the hair was yellowish brown, varying

from light brown to pure brown, and a coat of woolly hair, 2 to 214 cm.

in Jength, covered the whole body. Interspersed with these were a large

number of longer and thicker hairs which formed mane-like patches on

% Harte, Epovarp: “Les Mammiféres et oiseaux quaternaires connus jusqu‘ici en

Portugal. Memoire suivi d'une liste générale de ceux de la Péninsule Ibérique.” Com-

munic. du Service Géol. du Portugal, Tom. viii. pp. 22-85. pll. I-V. Lisbon, 1910.

*7 SALENSKY, W.: Uber die Hauptresultate der Erforschung des im Jahre 1901 am

Ufer der Beresowka entdeckten minnlichen Mammutcadavers.” C. R. Séa. Six. Congr.

Internat. Zoél.. pp. 67-86. Berne, 1904.

OSBORN, REVIEW OF THE PLEISTOCENE 305

the cheeks, on the chin, on the shoulders, flanks, avbdomen, ete. A broad

fringe of this long hair extended along the sides of the body as depicted

in the paleolithic sketches from the Combarelles Cave discovered by

Capitan and Breuil in 1901. Especially interesting is the food found

in the stomach and mouth, which consists of a meadow flora such as

characterizes this region of Siberia at the present day, thus appearing to

disprove the theory that the climate was milder than that now prevailing.

Nor does it appear that it was more frigid, because there are few repre-

Fic. 18.—The hairy mammoth (Elephas primigenius) and Paleolithic man (Homo

neanderthalensis )

Restored by Charles R. Knight under the direction of the author, 1914. Original in the

American Museum of Natural History, New York City.

sentatives of tundra vegetation. Grasses (Graminew) and sedges (Cy-

peracew) predominate. There were also wild thyme (Thymus), beans

of the wild oxytropis (Oxytropis compestris), seeds of the alpine poppy

(Papaver), and the boreal variety of the upright crowfoot (Ranunculus

acer), all still found in this region.

Woolly Rhinoceros—The woolly rhinoceros (D. antiquitatis, D. tich-

orhinus) is distinctly a cold-weather, or tundra form and the invariable

companion of the mammoth. Like D. mercku it has no front, or cut-

ting teeth hence it has been improperly considered as related to this spe-

cies, but it really belongs to the modern African group of Diceros (Ate-

306 ANNALS NEW YORK ACADEMY OF SCIENCES

lodus), which is distinguished by a very elongate front horn (Fig. 19)

and a small posterior horn, as in the existing white rhinoceros (D.

simus) of Africa. The resemblance of the cave drawings of the Pleisto-

cene animal to the lhving form is very close indeed except as regards the

heavy coat of hair, which, as in the mammoth, extends far below the

body. The hair of the face, of a golden brown color with an under-

covering of wool, is preserved in the St. Petersburg Museum. Through

a discovery (1911) at Starunia®* in East Galicia this animal is now

Fig. 19.—The woolly rhinoceros (Diceros antiquitatis)

Restored by Charles R. Knight under the direction of the author, 1914. Original in the

American Museum of Natural History, New York City.

completely known to us except the tail. The remains of the woolly

rhinoceros were found at a depth of 13.6m., including the head, the left

fore leg and the skin of the left side of the body, all with the muscula-

ture but lacking the hair. The Starunia specimen exhibits a broad,

truncated upper lip, small, oblique eyes, ears long, narrow and pointed,

a long nasal horn with oval base and shorter frontal horn, a short neck,

98 NIEZABITOWSKI, E. L.: “‘Die Uberreste des in Starunia in einer Erdwachsgrube mit

Haut und Weichteilen gefundenen Rhinoceros antiquitatis Blum. (tichorhinus Fisch.).

Vorlaufige Mitteilung.”’ Bull. Acad. Sci. Cracovie, Ser. B, pp. 240-267. April, 1911.

OSBORN, REVIEW OF THE PLEISTOCENE 307

on the back of which is a small hump quite independent of the skeleton.

The larger hump on the shoulders is formed by the long vertebral spines.

The legs are comparatively short. The skin is smooth. Niezabitowski

observes that D. antiquitatis resembles D. simus most closely, having in

common the elongate head with prominent orbits, the truncated upper

lip, the hump on the neck, and the short legs; it differs from D. simus

in the somewhat narrower muzzle, small, pointed ears and the presence

of a thick coating of hair.

Fic. 20.—Rhinoceros skulls

Skulls of the Pleistocene ‘“‘woolly rhinoceros,” Diceros antiquitatis of Eurasia (above),

and of the recent African ‘‘white rhinoceros,’ Diceros simus (below). In the American

Museum of Natural History.

Like D. simus, D. antiquitatis was a plains dweller living on grass and

small herbs. The woolly rhinoceros was confined more closely to the

edges of the great ice sheets than the mammoth: that is, it did not

migrate so far to the south, stopping at the Alps, while the mammoth

wandered into Italy as far south as Rome.

Elasmotheres—The elasmothere (Hlasmotherium sibericum) was an-

other companion of the mammoth which ranged over eastern Europe,

Germany, and southern Siberia. It was probably a steppe dweller. In

Pleistocene times it is reported as occurring as far south as Sicily.°° It

°° BranpT, J. F.: “‘Mittheilungen iiber die Gattung Hlasmotherium, besonders den

Schiidelbau derselben..” Mem. Acad. Imner. Sci. Pétersbourg, Ser. VII, Vol. XXVI, No. 6.

St. Petersburg. 1878. and Gaupry and Bove, “Matériaux pour l’Histoire des Temps

Quaternaires.” Fasc. 31. L’Elasmotherium. Paris, 1888.

308 ANNALS NEW YORK ACADEMY OF SCIENCES

is a gigantic animal, distinguished from all the European Pleistocene

rhinoceroses by the entire absence of the anterior horn and the pos-

session of an enormous horn situated on the forehead between the eyes;

also by the elaborate foldings of its dental enamel, to which the name

Elasmotherium refers. Its hypsodont and folded teeth were especially

adapted to a grassy diet, and Gaudry connects its appearance in Europe

with the extensive deforestation accompanying the Steppe and Tundra

periods of mammalian life. It apparently wandered into Europe from

central Asia and never became very abundant. The elasmothere is pos-

sibly descended?®° from the Aceratherium of the Upper Miocene of Ep-

pelsheim which has perfectly smooth, pointed nasals, and the rudiment

of a horn between the eyes.

Horses of the Pleistocene-——The horse was distributed all over the

northern hemisphere in Pleistocene times in the Glacial, Interglacial,

and Postglacial Epochs. In America no Postglacial horses are found.

In Europe horses were apparently abundant in Postglacial times and

two of the natural breeds appear to have given origin to two of the

modern domesticated types. The horses of the long warm Second

Interglacial Stage were remarkable for their great size (EF. siissenborn-

ensis, E. mosbachensis) which exceeded that of the largest recent breeds

(Pohlig, 1907).*° According to Pohlig the horses were at all times ac-

companied by the wild asses (H. hemionus) but this we are inclined to

believe was a special feature of the dry and cold steppe periods in which

we should expect to find asses similar to the dzeggetai of Asia of present

time. The existing wild ass. or kiang, of the Asiatic steppes certainly

appears in early Postglacial times at Wiidscheuer, Thaingen, Kessler-

loch, and Schweizershild associated with the late Aurignacian Palzo-

lithic culture. Reference of the ancient Pleistocene horses to Z. caballus

is certainly erroneous. The connection of these ancient Pleistocene

horses with the modern species and races requires further investigation.

We should expect to discover in Europe horses of three different habi-

tats or life zones, namely, of the dry African-Asiatic plams, of the

Eurasiatic forests and meadows, of the tundras and steppes. Such an-

ticipation appears to be verified through the new lines of study instituted

by Ewart?®? since 1904. Following more or less closely the work of pre-

vious students of the Equide he has shown that the different wild breeds

of horses have evolved in three kinds of environment. Thus we discover

10 OSBORN. HA. F.: “Frontal Horn on Aceratherium incisirum. Relation of the type

to Elasmotherium.” Science. N. S., Vol. IX, No. 214. pp. 161-162. Feb., 1899.

10 PoHLIG, H.: Eiszeit und Urgeschichte des Menschen. Leipzig, 1907.

12 Ewart. J. Cossar: “The Multiple Origin of Horses and Ponies.”” Trans. High-

land. Agric. Soc. of Scotland, pp. 1-39. 1904.

OSBORN, REVIEW OF THE PLEISTOCENE 209

horses adapted to: (1) forests and upland valleys; (2) high, dry, cold

steppes; (3) warm deserts and plateaus. In these three chief habitats

the horses may be respectively known as the “forest horse,” the “steppe

horse,” and the “desert horse.” Each has its distinctive coloring, tooth

structure, and proportions of the skull, body and limbs, in adaptation to

its peculiar mode of life.

The forest horse is relatively a large, clumsy animal. The face is

arched, as in the modern draught horse. The limbs are short, the front

cannon bone (Mic. III) being short and stout, the length only 514 times

the width. ‘he tail is depressed in contrast with that of the desert

horse. According to Ewart this type of horse (E£. robustus) occurs at

Solutré and in the Neolithic deposits of Ilford (Essex), and Kent. In

Aurignacean times Solutré was the site of a great open air Paleolithic

camp. Toussaint enumerates fragments of at least 100,000 horses,

which mingled with other bones of the chase formed a sort of rampart

around the camp. The majority of these horses belonged to the stout-

headed, short-limbed forest, or Norse type, measuring 54 inches (13.2

hands) at the withers, the size of the existing pony.’** The large joints

and hoofs were especially adapted to the low-lying marshy ground in the

- vicinity of forests, and the long teeth and powerful jaws were adapted to

feeding during parts of the year on coarse grasses, roots and other hard

substances. There is no evidence that the men of the Aurignacean age

either bred or reared these animals. The majority of the remains are

those of horses from five to seven years of age. This type of horse is

engraved on the walls of the cave of Combarelle, where the drawings are

chiefly of old Magdalenian age and the pure forest type of horse is most

frequently represented. ‘There is also a small, fine-headed type suggest-

ing the desert horse, and a larger, long-armed type suggesting the wild

ass.

The desert horse is the Pleistocene animal identified by Richard Owen

as an ass (#. asinus fossilis), but considered to be a horse by Ewart and

named by him £. gracilis. This is a small animal, not over 12.2 hands

in height, slender-limbed, with long, slender front cannon bones (Mtce.

III), the length being 714 times the width. The head is small, the face

fine and narrow, with a straight profile only slightly deflected upon the

cranium. The internal cusp (protocone) of the upper molars is short.

Remains of an animal of this type are found in the Pliocene of Italy

(small, slender-limbed varieties of H. stenonis) and France, and in the

Pleistocene of France and northern Africa. It agrees, so far as known,

with the existing Celtic pony type (FZ. caballus celticus), a variety of

103 WWART: Op. cit.

310 ANNALS NEW YORK ACADEMY OF SCIENCES

horse distinguished by small, fine head, large eyes, slender limbs, five

lumbar vertebree, now found in more or less pure form in the outlying

islands avd on the coast of western Europe. This animal is believed to

be a northern, hardy, thick-coated relative of the pure desert type, better

known as the Arabian, which gave rise to the modern thoroughbred.

Perfect representations of this type of horse are found in the engravings

and mural paintings of the Magdalenian artists in the caverns of Font

de Gaume, Combarelle, and Grotte de la Mairie.

A possible contributory to the desert breed of the Pleistocene and of

the modern domesticated horses is the animal of the E. sivalensis type of

the Upper Pliocene in the Siwaliks of India. This animal is tall, with

long, fairly slender limbs, long néck, well elevated tail, long face, which

is strongly deflected on the cranium with a slightly convex profile and

broad brow, and teeth with a narrow protocone.

Bears—The Postglacial bears (Ursus speleus) are found in greater

abundance than the lions. They include a gigantic and a smaller va-

riety. The former (Ursus speleus) nearly equalled the larger recent

bears in size and were more thick-set and of heavier proportions: the

front paws especially were of tremendous size. During a long period

the cave bears undoubtedly haunted the caverns undisturbed by Palzo-

lithic man and developed certain peculiarities of structure in these

haunts: thus the claw-bearing phalanges are feebly developed, indicating

that the claws had partly lost their prehensile function; the anterior

premolar teeth are practically vestigial: the cusps of the grinding teeth

are blunted in a way which is indicative of an omnivorous diet. It would

appear, therefore, that the large herbivorous mammals and even primi-

tive man found no very formidable enemy in the cave bear. While the

other and smaller races were contemporary, there are certain indications

that the smaller race (Ursus sub-speleus) was geologically older, bemg

found at Mosbach during the Second Interglacial Stage. Both races

became extinct during Postglacial times without leaving descendants.

The ancestor (Ursus arctos priscus) of the brown bear of Europe, by

some believed to be a descendant of the etruscan bear (Ursus etruscus)

of the First or Norfolkian Interglacial Stage, is also occasionally found

in the Pleistocene cave deposits. It is not so large as the cave bear and

while it has been mistakenly identified with the American grizzly (U.

horribilis) in reality it has closer affinities to the European brown bear

(Ursus arctos).

The badger (Meles tarus) also probably origimated in west-central

Asia, since the only other species known are confined to Asia. The two

OSBORN, REVIEW OF THE PLEISTOCENE 311

extinct Lower Pliocene species are found in Maragha, Persia (M. polakt,

M. Maraghanus) .*°*

TRANSITION TO THE EUROPEAN FOREST STAGE

This transition is believed to have begun late in Postglacial times,

toward the end of the Magdalenian culture period. Evidence that the

mammoth fauna lingered late both in the Dordogne region of central

France and to the north is found in the abundant representation of the

mammoths in the very latest paintings and engravings by the Magda-

lenian artists.

MIGRATION OF THE TUNDRA FAUNA

The backward or northward migration of the Tundra fauna is be-

heved to have occurred in the following manner.*® As the glacial caps

retreated they left barren stretches behind them and the valleys and

plateaus now free from ice became tundras where swamps alternated

with patches of polar willows and stunted fir trees, while other areas.

were covered merely with low, scrubby birches or reindeer moss and

- lichens. As these climatic conditions shifted northward before the re-

treat of the great Scandinavian glaciers the Tundra fauna followed. It

was a slow change that drove the Tundra mammals toward the dry re-

gions of the east to make room for the forests and their faunas advancing

from the south. It is clear that the north and east were the only direc-

tions of retreat for the damp climate and the spread of the woodlands.

RETREAT OF THE STEPPE FAUNA

As long ago as 1890 Nehring? held that the Steppe period of central

Europe was partly in Postglacial times. This opinion was supported by

Woldrich (1896),?°* and has been abundantly confirmed by Harlé’s ob-

servations in southern France and by the recent researches of Koken and

Schmidt. Steppe conditions of climate appear probable from the exten-

Sive depositions of the “Newer Loess” in Postglacial times (Koken,

104 ScHARFF, R. F.: The History of the European Fauna, p. 44. London, 1899.

10 STupER, T.: “Die Tierreste aus den pleistocenen Ablagerungen des Schweizers-

bildes bei Schaffhausen.’’ Neue Denkschr. allg. schweiz. Ges. Gesam. Naturwiss., Vol.

XXxv, pp. 1-38. 1896.

106 NEHRING, A.: Uber Tundren und Steppen der Jetzt- und Vorzeit, mit besonderer

Beriicksichtigung ihrer Fauna, pp. 81-166. Berlin, 1890.

107 WoOLDRICH, J. N.: “Ueber die Gliederung der anthropologischen Formationsgruppe

Mitteleuropas.” Sitzber. kgl. béhm. Ges., math. naturwiss. Class., 1896. Ref. Matiegka

in Centralblatt Anthrop., pp. 142-143. 1896.

342 ANNALS NEW YORK ACADEMY OF SCIENCES

Schmidt). On the other hand, Kobelt’®® and Scharff°® agree in think-

ing that the presence of Steppe mammals affords inadequate proof of the

steppe character of the country in Inter- and Postglacial times. The de-

posits of the “Newer Loess” in Postglacial times point to a dry steppe

period because according to the theory of Richthofen, which is now gen-

erally accepted, the loess owes its origin to wind-borne dust and sand

acting under the influence of a dry climate either in summer or winter.

The Steppe Fauna in deposits at several points is shown to have lin-

gered longer than the Tundra fauna. As regards the lingering of the

Steppe Fauna it is indicated in the succession of the three rodents char-

acteristic of the Tundra, Steppe and Forest conditions respectively,

namely :

Forest climate and conditions, the squirrel (Sciurus vulgaris)

Steppe climate and conditions, the jerboa (Alactaga jaculus)

Tundra climate and conditions, the banded lemming (Myodes torquatus)

The absence of fossil plants in the deposits of the steppe period is due

to the unfavorable conditions for the preservation of plant remains.

Small stretches of woodland were probably confined to the. banks of

rivers, to favorable mountain slopes, ete. The flora was probably like that

of eastern Hurasia or southwestern Siberia to-day. In their migrations

such animals as the jerboa which were unable to swim presumably a

the rivers while frozen over.

Sarga.—Of the Steppe fauna (fully described on p. 248) the saiga

antelope (Saiga tartarica) has at the present time retreated to the steppes

of eastern Europe and western Siberia. This animal is represented in the

carvings and engravings of Upper Paleolithic or late Magdalenian times

in the Dordogne region of France. Its fossil remains have been found in

thirteen localities in southwestern France in association with a cold

steppe fauna. In the same region have been found remains of the musk

ox (Ovibos).

SURVIVAL OF FOREST AND MEADOW FAUNA

The final retreat of the cold faunas of the tundras and steppes occurred

during the late stages of the Upper Paleolithic Magdalenian culture.

The most advanced Magdalenian art continues to represent the woolly

mammoth in the cavern of Font de Gaume (Dordogne) and elsewhere,

but in the very latest Magdalenian culture stages it would appear that

the mammoth and woolly rhinoceros were becoming rare. ‘This final

1088 KOBELT, W.: Die Verbreitung der Tierwelt. Gemissigte Zone. Leipzig, 1902.

109 ScHarrFr, R. F.: The History of the European Fauna. London, 1899.

OSBORN, REVIEW OF THE PLEISTOCENE 313

Magdalenian culture, which is correlated with the Gschnitz advance

(Schmidt, op. cit., p. 270), is later than the Steppe period of the “Upper

Rodent” layer, which is correlated with the preceding Bihl advance.

At the same time the Cro-Magnon, or Aurignacian type of Homo sa-

piens, which we believe to be the artistic race of the Reindeer period, dis-

appears or becomes greatly reduced in numbers and new brachycephalic

and dolichocephalc races of men enter Europe.

Azlan-Tardenoisian, Final Upper Paleolithic Culture.—This is re-

garded as the closing culture of Upper Paleolithic times. It is be-

lieved to be associated with the newly arriving broad-headed Furfooz-

Grenelle race. Although this point is not positively determined this

race is first found at Ofnet in Bavaria. It is readily distinguished from

the preceding Magdalenian culture by the degeneration of the stone in-

dustry into microlithic and other types and by the entire disappearance

of art in all its forms. The Azilian culture is essentially Paleolithic

although it embodies only its last degenerate stages. While the perfec-

tion of the older crafts was lost forever the Neolithic arts of polishing

stone, making pottery, cultivating land and domesticating animals are as

yet utterly unknown. The Azilian is the age of the stag for there is no

longer any trace of the reindeer or other Tundra forms. The bone imple-

ments are now made of the horns of the stag. The Tardenoisian culture,

supposed by some to be distinct from the Azilian, is characterized by flint

microliths of unusual fineness, but it appears that the Azilian and Tarde-

noisian cultures are contemporary (Obermaier, 1912).

There were two or more human races in Europe in these pre-Neolithic

times including brachycephalic and dolichocephalic types which are found

commingled at Furfooz. In the meantime Paleolithic races were ad-

vaneing in the north along the shores of the Baltic and preceding the

Campignian culture, which is the first of the Neolithic arrivals in the

Baltic region.

Forest Fauna.—The spread and multiplication of the Eurasiatic For-

est Fauna thus occurred before the close of Paleolithic times. Following

the retreat of the glaciers and the disappearance of steppe conditions of

climate there came a gradual subsidence of the coasts of northern Europe

and with it a more humid climate favorable to reforestation. Besides the

common squirrel (Sciurus vulgaris) which is the herald of forest condi-

tions all over the northern hemisphere, there appear in larger numbers

the entire Forest Fauna which we have traced from its beginnings in

early Pleistocene times and which we regard as having been resident in

favorable localities throughout the entire epoch. With the Tundra and

Steppe Faunas disappear also the wolverine (Gulo luscus) and the lion

314 ANNALS NEW YORK ACADEMY OF SCIENCES

(Felts leo spelea), which are never found in western Europe after the

Pleistocene although the lion lingered until a late period in eastern

Hurope.

The Alpine Fauna, which is mainly of central Asiatic rather than of

northern relationships, retreats to the higher levels of the Alps and the

Pyrenees. ‘Thus there remained in the forests, in the plains and in the

lower mountain regions of Hurope the direct descendants of the Hura-

siatic Forest and Meadow Fauna of the Pleistocene. It is noteworthy

that no new mammals appear in Europe at this time except those intro-

duced by man. The fauna of early Neolithic times is directly sequent

upon that of late Paleolithic times. This fauna has been discovered in

the Swiss lake dwellings’? (Fig. 9, 38-40) at Moosseedorf, Wauwyl,

Robenhausen, Concise, etc. In the peat bogs of Hassleben (41), etc., in

the travertines of Jena, Langensalza (42), etc.,** have been found the

following mammals:

Forest AND MEADOW

Bison bonasus, the European bison, still surviving in Lithuania.

Bos primigenius, collateral ancestor of the long-horned larger existing

eattle of western Europe. The “wrus,” of Cesar’s text. Surviving in

Germany until the sixteenth century.

Bos longifrons, the “Celtic short-horn,” the probable ancestor of the small

breeds of British short-horned and hornless cattle.

Cervus elaphus, the red deer or stag.

Cervus capreolus, the roe deer.

Alces machlis, the elk or moose.

Rangifer tarandus, the reindeer, surviving in central Europe until the

twelfth century.

Cervus dama, the fallow deer.

Sus scrofa ferus, the wild boar.

Sus scrofa palustris, the turf pig.

Equus caballus celticus, the Celtic pony, representative of the “plateau”

type.

Equus caballus typicus, the Norse, or “forest” horse.

Castor fiber, the beaver.

Sciurus vulgaris, the common squirrel.

Lepus timidus, the European hare.

Lepus variabilis, the arctic hare, in Ireland and the north.

Mus sylwaticus, the field mouse.

Arctomys marmotta, the marmot of the alpine fauna.

Ursus arctos, the brown bear.

Meles taxus, the badger.

100 RUTIMEYER, L.: “Die Fauna der Pfahlbauten der Schweiz,’ Neue Denkschr. allg.

schweiz. Gesell. gesam. Naturwiss., Vol. XIX. Ziirich, 1862.

ll PoHLiG, H.: “Vorliufige Mittheilungen iiber das Plistoczen, insbesondere Thiirin-

gens,” Sitzungsber. Niederrhein. Ges. Bonn, pp. 2-15. Mar. 3, 1884.

OSBORN, REVIEW OF THE PLEISTOCENE 315

Mustela martes, the pine marten, also the weasel, pole cat, the ermine, etc.

Lutra vulgaris, the otter.

Gulo luscus, the wolverine.

Canis lupus, the wolf.

Canis vulpes, the fox.

Felis catus, the wild cat.

ALPINE

Capra ibex, the ibex of the mountain or alpine fauna.

Rupicapra tragus, the chamois of the mountain fauna.

There is evidence of the “plateau” or “Celtic” horse in the Neolithic

deposits of Essex and of Switzerland (La Téne) ; it was widely distrib-

uted in Hurope and Asia in prehistoric times.*??

It is beyond the purpose of this volume to trace the history of domes-

tication. The dog (Canis familiaris), a descendant of the wolf (Canis

lupus), first appears in western Europe late in Upper Paleolithic

times.*4® The Neolithic immigrants, or men of the New Stone Age, pos-

sessed or brought with them cattle, sheep, goats, pigs, horses and dogs.

Appreciating the value of domestication, they certainly captured and

domesticated three indigenous European species, namely, the Celtic short-

horn cattle, the forest horse (#. caballus typicus) and the Celtic horse

(£. caballus celticus). The wild ox (Bos primigenius) was hunted but

not domesticated. The domestic ox (Bos taurus) shows many points of

resemblance to the Urus, but is not directly descended from it, but rather

from the Bos trochoceros type of the Pleistocene of Italy. Rutimeyer has

made an exhaustive study of this subject,*** tracing the origin of the

various types of domesticated cattle.

12 WWART, J. C.: Op. cit., 1907.

u8H), TROUESSART considers the diminutive wolf of India, Canis pallipes, as the prin-

cipal if not the sole source of all our races of domestic dog. This species of wolf, with

the exception of the Sumatran wolf, Canis sumatrensis, is also more closely related thin

any other to the dingo of Australia. “L’Origine préhistorique du chien domestique,”

Revue des Idées, pp. 388-411. June 15, 1911.

14 RUTIMEYER, L.: “Die Fauna der Pfahlbauten der Schweiz,’ Neue Denkschr. allg.

schweiz. Gesell. gesam. Naturwiss., Vol. XIX. Ziirich, 1862.

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ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXVI, pp. 317-383, pl. IV

Editor, EpMuNnpD Otis Hovey

PRESENT STATUS OF PHE PROBLEM OF

THE ORIGIN OF THE TETRAPODA

WITH SPECIAL REFERENCE TO THE SKULL

| AND PAIRED LIMBS |

WILLIAM K. GREGORY

meer ss NEW YORK

~ PUBLISHED BY THE ACADEMY

% Juxy, 1915

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[Annats N. Y. Acap. Scr., Vol. XXVI, pp. 317-383, Pl. 1V>~7-July, 19157

PRESENT STATUS OF THE PROBLEM OF THE ORIGIN OF

THE TETRAPODA, WITH SPECIAL REFERENCE TO

THE SKULL AND PATRED LIMBS?

By Wiii1am K. GREGORY

(Presented before the Academy, 13 February, 1911, 13 November, 1911,

11 December, 1911, and 10 February, 19138)

CONTENTS

. Page

Comparative study of the skulls of primitive fishes and Stegocephali..... 317

MMM MVIRURCSLNN AN Sco ae (ars Rael cin Svar egp od) Be o> aks ages ie 6 Res Wo chi cob aucihen, ei feketaie: oats Petes 317

eereaconerm:, Antiareht and vArthrodira.. 222. .3 0.6260 o.% eco kates es 318

oN eR SPUR EDP EN ORL LD Coes see co sree etch a Ghat Matas eG ia Wawa ae wae aD Cena See 320

IMU IR GET atany hy Sot Die eM ents SiiSed ays, UTI kw Re Powetay bk Bekok ee 321

MME eet 0 nt a. a le fac oe Say pte aynnad wi ote oo FG cel taienh GS moe en ew 322

POR RRAPEBE SE BM ohio at ete SOME Oe dace tae Me eid) a & jad on sin's ice oF eee wy Re Roernahe oe eee 325

eee SPE TE DL NR A as ee LNG occas & orca. goers 4 ah oganw GR aoe eo ant Bie A ee 334

Comparative study of the pectoral limbs of primitive fishes and Tetrapoda. 338

Origin and early history of the locomotive apparatus of vertebrates... 338

NE MDIMMIE SIDR E ert rer ryt es hi aM re ia, De Rae ee ye ui eee gd Mal 341

(SOE STE fod tel SENS MAMET ae BOLO Rte Dae = eae Ss on RPS aE ee RTS DRM 342

Pa PaDEREE BERKCEEDT PU eee sigh ee Tada a a hat as soy ia'ha ons rad or vceile 'aglal opaynore aw Mets Sok. pane 348

SMMRTERERM ERNE ae oy etd bP. Ones Se ew tin, coe laa Seite eGo te kmaia 345

RII Sen eh eet the daa a alata ated 2 a a TA ee eee ete ne EE De 348

NIE MRE SIOME es cette Rete hc IAL: SES AL SAI GH ch sug. dhe) aks bi ad gine Me Mee La ee ao 353

Pa SmCEO NIH icar ea cet eect aM east ti atone Masala poe te, ber acliea ake og a lave wernt 365

“Procoracoid and coracoid” or “coracoid and metacoracoid’’?........ 369

aM TED SARL Dre eal ia iNT tek Scie a ste ab wireke ie Sede, diese: Sac pantinn AON aa ace anes 375

na a Tiss Tice ila a gi Se ae Ae NR a eR SA a hs OF gare gh 376

COMPARATIVE STUDY OF THE SKULLS OF PRIMITIVE FISHES AND

STEGOCEPHALI

INTRODUCTION

The researches of Credner, Thévenin, Fritsch, Moodie and many others

on the amphibians of the Coal Measures, which are the oldest known

‘Tetrapoda, have served on the whole to bring into clearer focus the wide

structural hiatus that is still left between the known Amphibia and the

known fishes. For the former without exception have the paired limbs,

when present, in the form of cheiropterygia: while in the latter the

1 Manuscript received by the Editor, 6 March, 1915. (317)

= : —

Vis ie

the ta Oe

318 ANNALS NEW YORK ACADEMY OF SCIENCES

paired limbs are still incontestibly ichthyopterygia; and there are almost

equally great differences in the skull.

Baur’s view that the Amphibia and higher classes have been derived

from Paleozoic Crossopterygu has of late years been supported especially

by Watson (1912) and Broom (1913). Jaekel (1896. 1909), who de-

lights in such phylogenetic paradoxes, holds that the earhest Tetrapoda,

instead of having been derived from the fishes, have given rise to them

through progressive secondary adaptation to aquatic life. ,

Although the true linking forms between fishes and amphibia still

remain unknown and must be sought in formations of Lower Deyonian

or even Upper Silurian age (Moodie), comparative anatomy alone offers

sufficient evidence to prove that the recent fishes and Tetrapoda, in spite

of their admittedly wide differences and remote separation, yet trace their

origin to a common source. One need cite only the fundamental unity

of plan that runs throughout the vertebrate series and the weighty char-

acters that unite all the Craniata into a natural group. One recalls, for

example, that creatures so diverse in form and mode of life as fishes,

amphibians, reptiles, birds and mammals all have an identical arrange-

ment of the six eye-muscles in relation to the eyes and to three of the

cranial nerves; that the complex head is a synthesis of sensory capsules,

brain-trough, occipito-vertebral segments, primary jaws, branchial arches

and dermal elements; that the whole locomotive and muscular apparatus

of trunk, limbs and head is evolved out of metamerically arranged

myomeres of identical embryonic history. Thus, although the differences

between known Amphibia and known fishes are very marked, they are

morphologically of far less weight than the resemblances.

The main conclusion of the present paper is that among the various

groups of fishes which are known from the Devonian and Upper Silurian,

only one, the rhipidistian Crossopterygii, belongs near the ancestral line

or lines of the Tetrapoda. :

OSTRACODERMI, ANTIARCHI AND ARTHRODIRA

As the Silurian and Devonian Ostracodermi include the oldest known

types of chordate animals it is pertinent to inquire what their relations

may be to the remote ancestors of the Tetrapoda. All the known ostraco-

derms appear to be aberrantly specialized in certain directions, but long

consideration of their many peculiar characters has convinced me that

they stand far below the true fishes and that the group as a whole may

represent an important stage in the genealogy of the vertebrates.

Of the many beautifully preserved specimens of Birkenia, Lanarkia,

Drepanaspis, Pteraspis, Trematasms, Cephalaspis and allied genera, none

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 319

show the least indication of an internal skeleton, nor is there ever aly

trace of branchial arches and internal jaws. The pouch-like markings

on the inner side of the carapace in Pteraspis, the large sack-like mark-

ings in Thelodus pagei, the small round gill openings of several genera

indicate a branchial apparatus somewhat similar to that of the larval

Petromyzon. ‘The median opening in front of the paired eyes in Cepha-

lass and its allies suggests also the median nostril of Petromyzon. Due

caution is necessary in accepting any of these resemblances as indicative

of real affinity with cyclostomes, yet they apparently give evidence of the

mode of functioning of the mouth and pharynx.

In brief, the ostracoderms, instead of being free swimming predatory

types, with powerful locomotive organs, strong internal jaws and true

teeth, as in the later fishes and ancestral Tetrapoda, were, with the ex-

ception of the Birkeniide, sluggish and bottom-living forms accustomed

to feed on small organisms, which they obtained in the mud, or by draw-

ing water into their capacious pharnyx.

In these animals the process of cephalogenesis, or synthesis of elements

into a as head, was probably in a low stage of development, most

of the “cephalic buckler’” in forms like Drepanaspis and Cephalaspis

representing an expanded pharyngeal region rather than a large brain,

which may have been as small as it is in recent cyclostomes.

The ostracoderms represent a stage in chordate phylogeny immediately

preceding the acquisition of an endoskeleton impregnated with mineral

salts; their first visceral arches, if present, had not yet been transformed

into primary or cartilaginous jaws; the process of cephalogenesis was in

them in a low stage, and the elements of their shelly exoskeleton were

potentially homologous with the cosmine, vasodentine and isopedine of

primitive ganoids. ‘To that extent they stand in a pre-gnathostome stage

of evolution and probably represent the remote forerunners of the ganoids

and Tetrapoda, while possibly having remote relationships also in another

direction with the ancestral elasmobranchs.

The Antiarchi have advanced beyond the typical ostracoderms in hay-

ing the head sharply differentiated from the thorax and the mouth armed

with functional jaws, which are fashioned from the dermoskeleton. But

not even the excellently preserved specimens of Bothriolepis described by

Patten (1912) show any traces of car‘ilaginous jaws, branchial arches or

cartilaginous axial skeleton.

The systematic position and relationships of the Arthrodira constitute

so controversial a subject that it would be the part of discretion to avoid

it if-possible. Unfortunately this question as well as certain still larger

enes cannot be altogether kept out of the discussion as to the origin of

330 ANNALS NEW YORK ACADEMY OF SCIENCES

the Amphibia. With regard to the Arthrodira I can only say that I

have long pondered over the conflicting views and arguments of Smith

Woodward, Eastman, Dean, Hussakof, Tate Regan, Jaekel and Patten;

that I have endeavored to consider the question without bias of any kind

and have returned again and again to readjust the conflicting claims of

diverse evidence for the relationship of the Arthrodira: either with the

Dipnoi (Eastman, Smith Woodward), or with the Antiarchi (McCoy,

Dean, Hussakof), or with the Devonian Crossopterygu (Tate Regan) or

finally with the stem of the Dipnoi and Amphibia on the one side and

the Antiarchi, Ostracodermi and even the Arthropoda on the other

(Jaekel, Patten). Out of this maze of conflicting resemblances | gain

the impression that the resemblances which tend to brigade the Arthrodira

with the Antiarchi are quite specific and of evidential value,? while the

resemblances in the jaws and head-shield to those of Dipnoi and still

more to those of Crossopterygii, are, with certain exceptions, of a vague

and inconclusive character, suggestive of convergent evolution between

widely removed groups. And in this connection I would call attention

to the arthrodiran appearance of the skull-top of the modern armored

catfish Claritas magur (figured by Goodrich, 1909, p. 381), as illustrating

the deceptive effects of convergent resemblances; a similar example is the

Cephalasprs-like appearance of Loricaria.

Until it shall be learned whether Macropetalichthys is really related

to the Arthrodira and what its affinities are with other groups, it is

hardly likely that the Arthrodiva problem can be definitely settled. In

the meantime I must hold provisionally: first, that the arthrodiran jaws,

skull and thorax represent progressive, carnivorous adaptations from a

generalized antiarchian stem ; secondly, that the resemblances in the skull-

roof to those of Dipnoi and Crossopterygil have been independently as-

sumed; thirdly, that the Arthrodira are not true gnathostomes and con-

sequently not anywhere near the line leading toward the Tetrapoda, but

that they are, on the contrary, the last and highest of a pre-gnathostome

adaptive radiation. In brief this pregnathostome radiation was char-

acterized by an accelerated evolution of the exoskeleton of the head and

thorax and by a retarded evolution not only of the endoskeleton but of

the whole locomotive apparatus.

ELASMOBRANCHII

In the Acanthodii, which are the earliest elasmobranchs, the gnatho-

stome series is first made known by its relatively advanced adaptations

to predatory, free swimming habits: first, the whole pharyngeal region

> Cf. the list of common characters given by Hussakof. 1906, pp. 123-136.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 321

is supported by a stout cartilaginous framework and the contractility of

the anterior pair of arches has led to their functioning in the seizing

and ingestion of the prey; secondly, the primary locomotive organs,

namely, the myomeres, have much increased in number and size, the head

becoming relatively small; thirdly, the paired sensory capsules (olfactory,

optic, auditory) and the primitive brain-trough are synthetized into a

chondrocranium, which very early affords support for the oral arches,

while the more active life necessitates the strengthening of the primitive

endoskeleton, an end which is attained in this group by the deposition of

calcareous salts in the hyaline cartilage; fourthly, many accessory struc-

tures appear: such as median and paired fins and fin supports, fin-spines,

true teeth and minute rhombic scales.

All these external improvements in the locomotive and food-getting

structures were no doubt matched by corresponding increase in the size

of the brain and probably by marked changes in the proportions and

placement of its principal parts.

Such a transformation from pre-gnathostome to primitive gnathostome

conditions was a critical step in the history of the vertebrates and makes

the differences between Tetrapoda and true fish seem relatively unim-

portant.

The elasmobranchs are excluded from direct ancestry to the Tetrapoda

by the fact that they too early overspecialized in some respects while

remaining on a low stage in others: first, the exoskeleton was formed too

largely from the outer layers of the many-layered skin, the deeper, strati-

fied connective tissue remaining unossified ; hence except in the Acathodii

we see an armature of thorny denticles or placoid tubercles rather than

of osseous plates and scales. And secondly, the endoskeleton, instead of

hecoming osseous, became thoroughly calcified.

The earliest elasmobranchs have the preorbital portion of the cranium

short, the small eyes being almost terminal and not widely separated from

each other, as in most of the ostracoderms, antiarchs, arthrodires, earliest

ganoids and tetrapods. It is only in specialized types in all these groups

that the olfactory capsule or rostrum grows forward and the eyes are

displaced backward and become of large size.

So far as known the elasmobranchs lack the median opening between

the orbits which is doubtfully homologized with the pineal opening in

ostracoderms, antiarchs, arthrodires and primitive ganoids.

ACTINOPTERYGII

The Dipnoi, Crossopterygii and Actinopterygii (collectively known as

Osteichthyes) share with the elasmobranchs on the one hand and with

BOO ANNALS NEW YORK ACADEMY OF SCIENCES

the tetrapods on the other all the characters which separate the Gnatho-

stomata from the Agnatha. But they have advanced bevond the elasmo-

branchs toward the tetrapod type in many particulars, especially: (1) in

the appearance of true bone cells both in the exo- and endoskeletons; (2)

in the formation of scales, dermal rays and dermal plates, having in the

most primitive forms an outer cosmine layer, a middle zone of vasodentine

and basal layers of isopedine; (3) the formerly continuous dermal cover-

ing of the head and trunk is now fragmented into the dermal plates of

the skull, of the branchial region (operculars, gulars, etc.) and of the

pectoral region; (4) the spiracular cleft is usually closed.

Most of the known Osteichthyes are excluded from the ancestry of the

Tetrapoda by various specializations, either of the exoskeleton or of the

median and paired fins as noted below.

In the Actinopterygii, a sufficiently full morphological series enables

us to follow the changes in the group, beginning with the very generalized

Deyonian Chetrolepis and culminating in the most specialized types of

modern teleosts. In the most primitive of the series (Paleoniscide) the

elements of the skull-roof may be in general homologized with those of

the most primitive Crossopterygul, although the details are quite ditferent ;

there is, however, no special resemblance to the skull of the earliest

Stegocephah, except in so far as the elements of the skull-roof include

paired frontals, parietals, pterotics and other paired elements. ‘The brain

also is diversely specialized in the surviving forms.

DIPNOE

A closer structural approach to the tetrapod type is attained in this

group. In the brain the Dipnoi have retained the well-developed olfac-

tory lobes and cerebra, which are requisite for the ancestral tetrapod.

‘hey also have a functional lung and, as Nellicott (1905) has shown,

the venous system presents close ontogenetic resemblances to the urodele

type. The larve and embryos of the Dipnoi, as well as of the Crossop-

terygil, reveal further well-known striking resemblances to the urodeles,

and, as noted below, there are many features of the locomotive organs

and mode of locomotion in Dipnoi that foreshadow the conditions in

VYetrapoda. While much of this might be ascribed to convergence, it all

imphes a similarity in the “potential of evolution,” that is, of structural

possibilities, in the forerunners of these groups.

The known Dipnoi are all excluded from direct ancestry to the

Amphibia by the specialized character of the dentition, including the

formation of complex radially arranged tritoral plates on the roof of

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 323

the mouth and on the inner side of the mandible and the loss of marginal

teeth on the premaxille, maxille and dentaries.

The earliest Dipnoi have “paralleled” the Tetrapoda and other pro-

gressive types in that the preorbital rostrum is expanded and the orbits

are relatively far posterior though still of small size; the pattern of the

skull-roof, with its paired “frontals,” “parietals” and other elements, is

also probably analogous, rather than homologous, with that of Tetrapoda.

Our conception of the relationship of the Dipnoi to the Tetrapoda is

Fic. 1.—Pattern of skull-top of Devonian dipnoans

A, Dipterus, after Goodrich, slightly modified; B, Scaumenacia curta, after Hussakof.

In specimens of Dipterus the numerous sensory pits are scattered over broad tracts,

the general directions of which are indicated by the dotted lines, except in the occipital

region where the dotted lines represent shallow grooves. The principal sensory tracts

are in general similar to those of Stegocephali.

The “‘parietals,’’ “frontals,”’ etc., are probably analogous rather than homogenous with

those of Tetrapoda.

Dso, dermosupraoccipital ; S.¢, supratemporal (pterotic) ; 7b, tabulare (epiotic) ; P.p,

preparietal; Fr, frontals; Na, naso-ethmoid region.

to some extent dependent upon the validity of Dollo’s view (1895) that

Dipterus is the most primitive known Dipnoan, structurally ancestral to

all the later types. The skull top of the modern Ceratodus is so widely

different from that of Dipterus that Dr. Eastman (1907, p. 95) has con-

cluded that the two forms have no near relationships with each other and

that the living genus represents some other and independent line that

has come down from a pre-dipterine stock. But after comparing the

skull patterns of Ceratodus, Ctenodus, Phaneropleuron and Scaumenacia

394 ANNALS NEW YORK ACADEMY OF SCIENCES

T incline to the opinion that Dollo’s view is the correct one. The skull

pattern of Scaumenacia, as figured by Hussakof (1912, p. 136), is suffi-

ciently close to that of Dipterus ( Pander, 1838, Tab. 3) to enable one to

homologize the principal bony elements in these two genera (Fig. 1).

The Scaumenacia skull, I think, points the way to that of Phaneropleuron

(Goodrich, 1909, p. 239) and this in turn brings us within reach of the

Ceratodus skull-pattern. In short it appears probable that the Ceratodus

skull-pattern has been derived from a Dipterus-like type through the

following changes: (a) the small frontals and parietals have become

enlarged, so as to cover the median, pre-parietal element; (6) the oppo-

site frontals have united to form the wrongly called “ethmoid” of Cera-

todus; (c) the opposite parietals have united with each other and with

the median supraoccipital or nuchal plate to form the large median

“occipital”; (d) the longitudinal series of small elements running from

above the orbits to the back of the skull have coalesced and broadened,

to form the large pair on either side of the median plate; (e) the small

plates behind the orbit and above the operculum have coalesced and

broadened into the large plate marked pterotic in Goodrich’s Fig. 206

(1909, p. 237); (f) the remaining small elements immediately behind

the orbit are part of the circumorbital series: (g) the exposure of the

chondrocranium above the occiput and above the front part of the skull

T regard as quite secondary and as a relatively modern specialization;

the massive character of the chondrocranium may be due to the fact that

the stout tritoral plates on the roof of the mouth and the heavy mandible

require a massive firm skull for their support: it may also be related with

the fact that the scales and dermbones of the skull are losing their dense

osseous character and becoming horny. The exposure of the chondro-

cranium is carried to an extreme in Lepidosiren, and it cannot be claimed

that this degenerate, eel-like form is more primitive than the numerous.

Palzozoic Dipnoi, Crossopterygii and Actinopterygii that have a complete

skull-roof. iar ao

In view of the foregoing discussion it is hardly necessary to state that

I am unable to accept Dr. Eastman’s comparison (1907, p. 132) of the

Ceratodus skull with that of Dinichthys at its face value, and, with Dr.

Dean (1907), I must ascribe these resemblances to convergence, referring

again to the arthrodiran appearance of the skull-top of one of the modern

catfishes (vide supra, p. 320).

Returning to Dipterus I would also be inclined to disagree with Good-

rich’s statement (1909, p. 256) that “the arrangement of the cranial

covering bones points rather to the dipterids being a highly specialized

offshoot from the base of the dipnoan stem.” As already stated, the

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 325

Dipterus skull-top certainly includes the same elements as those of Scau-

menacia, the chief difference being that these elements are smaller and

that there are a number of other small elements in the supraorbital-

occipital series, which may later have united with adjacent elements.

The Dipterus skull-top also makes a distant approach to the primitive

rhipidistian type seen in Osteolepis, in so far as it retains lines of sensory

pits running longitudinally above the orbits and extending back to the

transverse nuchal commissure; both skulls also have a circumorbital

series, a supraorbital-occipital series, a short rostrum, paired frontals,

parietals and other paired elements, the chief difference being that in

Dipterus there is a prominent median preparietal and that the nuchal

plates are more closely united with the occiput.

- The conclusion that Dipterus is the most primitive of all the Dipnoi

is further supported by the remarkable agreement in histological struc-

ture of the dermoskeleton of Dipterus and that of the crossopterygian

Osteolepis as shown in Pander’s superb sections (1858, Tab. 5, Figs. 1,

17; 1860, Tab. 5, Figs. 1-8, 22). In both cases the dermocranium con-

sists of a thin outer layer of ganoine, which covers the cosmine layer,

underneath this is a vascular layer and on the bottom a stratified hori-

zontal layer. This histological pattern is also repeated in the scales of

both Dipterus and Osteolepis, and:to my mind is of great weight in indi-

cating the common origin of the Dipnoi and Crossopterygu. Add to that

the facts: that in both groups the earliest members have the paired fins

of the mesorhachic or biserial type; that both have two dorsal fins; that

both have a heterocercal tail provided with similar dermal rays, and we

have a case for the common origin of the Crossopterygii and Dipnoi,

which is further strengthened by the well known resemblances between

the modern Polypterus and the dipnoans in the early stages of develop-

ment. Nor should the differences in brain structure of these modern

forms outweigh the above mentioned resemblance, for there is no evidence

that the brains of the Devonian Crossopterygu and Dipnoi were any more

divergent from each other than were the other parts of the body.

CROSSOPTERYGII

The known Dipnoi being excluded from direct ancestry to the Tetra-

poda by reason of certain specializations of the skull and dentition, what

ean be said of the claims of the Crossopterygii ?

_Polypterus, and doubtless also its near ally Calamoichthys, which are

the only surviving crossopterygians, have become highly ichthyized in

brain characters and thus are far removed from both the dipnoan and

the amphibian types; but as stated above it may well be that the Devonian

296 ANNAES NEW VORK ACADEMY OF SCIENCES

Crossopterygu had a more primitive brain structure. In Polypterus the

scales and dermal plates have become like those of the primitive Actinop-

terygu (Goodrich 1909, p. 292), the cosmine layer being covered exter-

nally by stratified ganoine layers; but for several reasons I regard this

condition as secondary. Nor can I accept Goodrich’s view (1909, p. 300)

that Polypterus should be removed altogether from the Crossopterygii.

In any event it.is further removed from relationship with the Tetrapoda

than were its Devonian predecessors.

The only crossopterygians that can claim even remote relationships

with the Amphibia are the Devonian Rhipidistia, especially the Osteole-

pide and the nearly allied Rhizodontide. Dr. A. 8S. Woodward’s com-

parison (1898, pp. 24, 25) of the skull-roof of Rhizodopsis, representing

the Rhizodontide, with that of Pelosauwrus, representing the Stegoce-

phalia, serves, however, to emphasize the remoteness of this relationship

and to raise the question whether the supposedly homologous elements,

as the frontals, parietals etc., in the two groups, may not after all be

analogous rather than truly homogenous. Dr. Moodie’s comparison

(1908) of the sensory canals of the skull-rocf of stegocephalians with

those of Amia and Polypterus offered some reimforcing testimony, which

would have been strengthened if Pander’s superb figures showing the sen-

sory pits and skull elements of Osteolepis, Diplopterus, Dipterus and

other genera had been taken into consideration. Watson’s comparisons

(1912) of the skull-base of Rhizodus with that of certain Carboniferous

stegocephalians (Pteroplax etc.) having a single median occipital condyle

and a continuous or non-fenestrated palate, further strengthened the case.

A wider basis of comparison appearing desirable, 1 have made during

the last few years repeated comparison of the skull patterns of various

Paleozoic and recent fishes with each other and with those of stegoce-

phalians of all known orders. Original material of Osteolepis, Megalich-

thys, Dipterus and of many other fossil and recent Osteichthyes has been

studied, and for the skull patterns of many of the Palaeozoic fishes I have

also had recourse to the excellent figures of Pander, Huxley, Traquair,

Smith Woodward, Goodrich, Wellburn (Megalichthys), Jaekel (Dviplop-

terus) and others. Similarly among the Stegocephali the American

Museum collections have furnished for study typical examples of the

principal groups; and in making comparisons with the fishes I have also

had before me the figures of Credner, Fritsch, Moodie, Fraas, Embleton

and Atthey (Loxomma), Watson, Broom, Williston and others.

Before considering in detail the skull-roof in primitive fishes and Tet-

rapoda, it may be appropriate to ask whether any general adaptational

reasons can be assigned provisionally for the several characteristic ar-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 327

rangements of the sutures which produce the distinctive skull patterns

of these early types. It was recognized by Ryder (1892) that the shape

and arrangement of the scales of fishes was originally a result of the moye-

ments of the myomeres, but so far as J know it has not hitherto been

taught that the shape and arrangements of the dermocranial elements,

which in primitive fishes are histologically homologous with scales, were

likewise the result in part of muscular strains and stresses. It may be

stated as a general hypothesis that in the dermocranium of primitive

fishes the position and arrangement of the sutures and the consequent

pattern of the osseous “elements” are the evolutionary resultants of the

various symmetrically balanced stresses induced by the action of the

underlying muscles of the eyes, jaws, branciial arches and pectoral limbs.

mm composition with the position and size of the olfactory, optic and

auditory capsules. It ts at least a fact that sutures and other articula-

tions define loci of relative mobility, centers of ossification define loci of

relatwe stability. Differential growth of one region of the skull, as in the

rapid elongation of the snout, also results in more or less rearrangement

of the sutures and osseous elements.

I would also advance the hypothesis that the whole dermoskeleton of

the head in the unknown pre-Devonian forerunners of the Dipnoi, Cros-

sopterygu and Actinopterygii was a continuous membranous covering,

consisting (see Goodrich, 1909, pp. 215-217) of minute cosmine tuber-

cles underlain by imperfectly differentiated vasodentine and isopedine

and entirely homologous with the membranous covering of the trunk and

fin-folds. Wherever movements of this membranous covering were pro-

nounced, as in the opercular region, around the orbits and near the spi-

racular clefts, there were deep creases, but where movements were less

pronounced the creases were proportionally shallow. When the critical

stage of ossification began, in the forerunners of the Dipnoi, Crossop-

terygu, Actinopterygu, the roof of the dermocranium commenced inde-

pendently in the several lines a process of fragmentation; so that the

rostrofrontal segment was split up into premaxille, nasals, ethmoids,

frontals; and the orbito-parietal segment was divided into its component

“elements.”

The excessive subdivision of the maxillary and preoperculum of Lej-

dosteus, of the parafrontals of ccelacanths, of the spiracular plates of

Polypterus furnish extreme examples of fragmentation. In the skull-

root of Osteolepis, on the other hand, we may have an example of a form

in which ossification of the dermecranium had already taken place, but

fragmentation of the “cranial buckler” was still incomplete. In some

specimens of Osteolepis the median suture between the frontals extends

328 ANNALS NEW YORK ACADEMY OF SCIENCES

forward nearly to the tip of the rostrum; in others it is much shorter;

none have the separate nasals which appear in higher types; on the other

hand, the transverse occipital row of small plates (see below, p. 329) seen

in Osteolepis and Gyroptychius, are barely indicated in Tristichopterus

and had not split off in Rhizodopsis or Megalichthys.

As the conical teeth of Crossopterygii were on the outer margins of the

jaws and the movements of the jaws were chiefly vertical, the suggestion

may perhaps be hazarded that the transverse stresses conditioned the

evolution of the longitudinal median sutures, while the anteroposterior

stresses resulted in transverse sutures, those separating the rostro-frontal

and postorbito-parietal segments being in Osteoleyis particularly deep.

At any rate the resulting skull pattern includes many more or less rect-

angular plates, as also in the Actinopterygu. In the primitive Dipnoi,

on the other hand, the teeth were clustered in fan-like ridges on the roof

of the mouth and the movements of the jaw were more oblique; this

possibly may have partly conditioned the oblique character of the skull-

roof sutures and the polygonal form of the elements. In the Stegocephali

both rectangular (e. g., Branchiosaurus) and oblique (e. g., Stegops

dwaricata Moodie) types occur.

After the primary sutures had been established and the continuous

dermocranium fragmented into small plates, there usually followed an

enlargement of certain plates, and sometimes a coalescence of adjacent

plates, as in the Dipnoi and other groups of fishes; such enlargement and

coalescence following ex hypothese not only upon the shifting of lines of

greatest mobility or the diminution of mobility between elements, but

also from the circumstance that as a general rule in evolutionary series,

after individual structures have been differentiated out of a former con-

tinuum they appear to acquire a certain evolutionary initiative, so that

they may enlarge or decrease or shift their position, or crowd apart their

fellows in a manner strongly hereditary, but often without any assignable,

immediately adaptive purpose.

The position and apparent homologies of the elements of the skull-

roof in the Rhipidistia, in comparison with other primitive fishes and

with the Stegocephah, are as follows:

The orbits in primitive Crossopterygii are small and sometimes far

forward (Osteolepis, Fig. 2), quite near the front of the nasal rostrum,

which is extremely short and obtuse; the same is also true of the most

primitive Actinopterygu (Paleoniscide). The roof of the nasal rostrum

(ethmoid) is undivided and there are no paired nasal bones. The short

frontals, lying between the orbits, almost form part of the rostrum.

Between them in some genera (Osteolems, Glyptopomus, Diplopterus)

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 329

lies a foramen opening into a median tube (Pander, 1860, Tab. 5, Figs.

1-5) which has been identified by authors (e. g., Newberry, Dean, Jaekel)

as the pineal foramen. In Dipterus also the frontals are short, but in the

Paleoniscide they are longer. The postfrontals in Osteolepide and

Rhizodontide are elongate narrow elements extending from the postero-

superior margin of the orbits to the pteroties (“squamosals”).

The parietals in the Rhipidistia are longer than the frontals, from

which they are separated by sharp transverse sutures. In Dipierus, on

the other hand, the parietals were short, in Paleoniscide they were short

and wide.

_ Behind the parietals in Osteolepis microlepidotus (Pander, 1860, Tab.

I) was a transverse series of small quadrangular elements, closely ap-

pressed to the parietals and “pterotics,” which at first sight suggest the

transverse occipital series of Stegocephali. The innermost or median

pair of this series suggests the paired dermosupraoccipitals or postparietals

of Stegocephali; next to these comes a second or middle pair; the third

pair at first suggest the tabularia, and the outermost pair in the species

under consideration suggest the “epiotic cornua’ > of such Stegocephali as

Archegosaurus.

Nevertheless, in spite of these attractive resemblances with the trans-

verse postparietal series in Stegocephali, I am finally constrained to treat

them as analogies only. First, this transverse postparietal series is best

developed in Osteolepis microlepidotus, is imperfectly developed in O.

macrolepidotus, Gyroptychius, Tristichopterus, and is entirely absent in

Rhizodopsis, Megalichthys, Glyptopomus and Holoptychius. But struc-

tures that are developed only in a few forms, rather than in whole groups,

cffer very unsafe guides for homologization with similar elements in other

widely separated groups, a principle too often neglected by comparative

anatomists. Secondly, this transverse occipital series in Osteolepis does

not bear a transverse line of sensory pits, as it should do if it were

homologous with the transverse postparietal series of Stegocephali (cf.,

Moodie, 1908). Thirdly, there is another row of transverse plates consti-

tuting the nuchal or “supratemporal series,” lying behind the occipital

suture, which bear a transverse sensory line and are homologized by all

authorities with the postparietal row in Stegocephali. This nuchal series

_ may terminate dorsally either in a single median plate, as in Dipnoi, all

Rhipidistia, many Actinopterygii (?supraoccipital), or in double or

paired median plates, as in Polypterus, Amia, Lepidosteus and all Stego-

cephali.

Concerning the nuchal series in T'ristichopterus alatus, one of the

Rhizodontide, Traquair (1875, p. 386) wrote as follows:

330 ANNALS NEW YORK ACADEMY OF SCIENCES

Fig. 2—Skuli patiern of Osteoclepis microlepidotus

A primitive Devonian rhipidistian.

After Pander, but with revised identification of elements.

A, Top view. B, Side view.

Supposed homologies with the skull elements of Tetrapoda are indicated by the ab

breviations.

Dorsal series: Na, naso-ethmoid region: F. frontal region with median foramen; P#-f.,

postirontals ; Pa, parietals; behind the parietals are the parieto-occipital series, perhaps

not differentiated in Tetrapoda, as they lack the occipital commissure of sensory pits

(see text).

Circumorbital series: P.f, prefrontal; La, lacrymal: Ju, jugal (suborbital) ;-Po.o, post

orbital.

oe series: S.f, supratemporal (piterotic) : Sg. squamosal (preoperculum, cheek

plate ““X"*). :

Gnathal series: Mr, maxilla (the premaxilla is not separated from the naso-ethmoid

mass); Q.j, quadratojugal (anterior interoperculum. cheek plate “X’’); D, dentary;

Sp, splenial (anterior infradentary) ; Ang, angular (middle infradentaries) .

Opercular series (lost in Tetrapoda) : Op, operculum; S.op, suboperculum; Gu, gular;

T.op, interoperculum ( ?may have fused with quadratojugal in Tetrapoda).

Nuchal-occipital series: Dso, dermosupraoccipital (postparietal. median supratemporal) ;

Tb. tabulare (epiotic. lateral supratemporal). =.

oS eV"

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 331

“Along the posterior margin of the cranial shield are three plates (s. f,

figs. 1 and 2), one mesial, somewhat polygonal in form, and two lateral, each

apparently of a triangular shape. These are obviously the representatives of

the three plates, which occur in a similar position in Osteolepis, Glyptolemus,

Megalichthys, &c., and of which different interpretations have been given by

different authors. In Professor Huxley’s description of Glyptolemus,? the

mesial one is marked ‘supra-occipital,’ the two lateral ‘epiotic. Mr. Parker ’

has, however, pointed out that they are dermal bones, and not to be considered

homologous with those other deeper ossifications of the cranial cartilage. By

Pander’ they are in Osteolepis simply designated ‘Hautknochen’ and consid-

ered to be equivalent to the five little plates, which in the recent Polypterus

occur immediately behind the transverse row of supra-temporals, and between

the pair of upper supra-claviculars (supra-scapulars), being in reality the first

scales of the back. On the other hand, he considered the transverse chain of

small plates (supra-temporal) which lie immediately behind the parietals of

Polypterus, to be representel in Osteolepis microlepidotus by the narrow por-

tion of the cranial shield, which in that species is marked off near the hinder

margin by a more or iess interrupted superficial transverse groove. I am my-

self very much inclined to the belief that the three dermal bones in question

are in reality equivalent to the transverse supra-temporal chain in Polypterus

and Lepidosteus, and which have their representatives as well in the am-

phibian Labyrinthodonta as in most Teleostean fishes; the transverse grooving

across the posterior part of the cranial shield in many Saurodipterines being

probably only of the nature of superficial markings.”

In brief, the skull-roof of Rhipidistia may be regarded as consisting of

the following regions or segments: first a rostrofrontal segment, com-

prising the parts which later became separated into the premaxille,

ethmoid, nasals, frontals and postfrontals. In Osteolepis this is sep-

arated from the next segment by a sharply defined transverse suture.

‘he second or parietal segment of the skull-roof includes only the elon-

gate parietals and the wrongly named “squamosals’ (pterotics). The

third or occipital segment which is rarely distinct from the preceding

(Osteolepis, etc.) consists of a transverse row of small elements, which,

in spite of their resemblance to the dermoccipitals and tabularia of Stego- »

cephali, are probably not homologous with them. The fourth or nuchal

segment, which originally pertained to the pectoral girdle rather than to

the skull, is sharply separated from the true occipital segment; it usually

consists of a single median and a pair of lateral elements; all three of

these are usually named “supratemporal,” but they are probably homol-

_ ogous with the paired dermosupraoccipitals and tabularia of Stegocephali.

Returning to the region above and behind the orbits we find not only

3 Dec. Geol. Survey, x. p. 2.

* Shoulder Girdle and Sternum, p. 19.

5 Ueber die Saurodipterinen, &c., des Devonischen Systems, p. 11-12. St. Petersburg,

1860.

339 ANNALS NEW YORK ACADEMY OF SCIENCES

in primitive Rhipidistia but also in Dipterus (Fig. 1, 4) a longitudinal

chain of small elements, bearing the supraorbital sensory line and extend-

ing back to the occiput. This series corresponds in position with the

postfrontals, intertemporals and suprasquamosals (pterotics or true supra-

temporals) of Stegocephali.

The circumorbital series of Rhipidistia and primitive Dipnoi while

variable in number and form likewise correspond in position, and appear

to be collectively homologous with, the prefrontal, lacrymal, jugal, post-

orbital and postfrontal of Stegocephali.

The spiracular cleft, which in Holoptychius, Osteolepis microlepidotus

and Polypterus as well as in Dapedius is located behind and above the

orbit, appears to be lost in many Crossopterygii and in all Dipnoi, as

well as in the Stegocephah.

The elements of the opercular series of Osteolepis and other Rhipidistia

eppear to be only partly homologizable with those of Dipterus (Goodrich,

1909, p. 240) ; in the Stegocephali this series appears to have been lost

in part and in part preserved. In Osteolepis the preoperculum plus cheek

plate (X) was external to the quadrate ; it was also behind the postorbital

and jugal, and below the pterotic or true supratemporal; in all these

respects it corresponds perfectly with the lower temporal bone of Tetra-

poda, which is by many authors identified with the mammalian squa-

mosal. In Osteolepis and Diplopterus the preoperculum also bears a

sensory tract, as does also the squamosal of certain Stegocephali (e. g.,

Ceraterpeton, Tutidanus, Moodie, 1908). In other Rhipidistia (e. g.,

Tristichopterus, Diplopterus, Megalichthys) the preoperculum has an

essentially identical position, but includes two elements, a posterior one

corresponding to the back part of the preoperculum of Polupterus and

to the whole of the preoperculum of Amia and an anterior one (X)

corresponding to the posterior row of postorbitals in primitive Holostei,

as suggested by Traquair.

Below the preoperculum (squamosal) in Osteolepis mnicroleaaeae

Diplopterus, Megalichthys, etc., are one or two elements (X’), together

corresponding in position to the interoperculum of higher fishes, which

have the topographic relations of the quadratojugal of Stegocephali and

like that element also bear a sensory tract.

The retention and full development of the operculum, suboperculum,

median paired and lateral gulars sharply separate the Rhipidistia from

the Stegocephali. The fate of these elements is discussed below (p. 337).

The under side of the skull should, on general principles, yield impor-

tant evidence bearing on the problem of the relationships of the Tetra-

poda and the Rhipidistia. Unfortunately this region was not described

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 333

by Pander or Traquair and seems to be known only in Megalichthys. In

this genus, Watson (1912, pp. 9-10) has noted the following remark-

able resemblances to the very primitive Stegocephalians Pteroplaxz and

Loxzomma:

“The Basisphenoid of Megalichthys has sometimes carotid foramina just as

in Lozomma. It has small but distinct basi-pterygoid processes which are,

however, not provided with articulating surfaces but with sutural ones. The

long parasphenoid extends forward to the premaxille as it may do in Ptero-

plax. Its lateral borders are in contact with the Pterygoids, to which they

afford support, and the bone is connected with the roof of the skull by a fused

ethmoid.

“The Pre-vomer is identical with that of ‘Lozomma’ in the majority of its

attachments, carries one large tusk and a pit for the replacing tooth. It meets

its fellow of the opposite side, and forms the front of the posterior naris; it is

doubtful, however, if it meets the palatopterygoid.

“The Palatopterygoid of Megalichthys is exceedingly like the palatine and

pterygoid of Pteroplax. They have similar relations to the basisphenoid,

parasphenoid and maxilla. There is the same row of small teeth parallel to

those of the maxilla with larger teeth inside them, and the.pterygoid is cov-

ered with the same shagreen of fine teeth.”

Watson also states that unlike later Stegocephali Loxomma has a single

median occipital condyle on the basioccipital, and that this condyle “ex-

actly resembles the end of a vertebral centrum, which it no doubt is.”

In these features Loxomma therefore agrees with Rhipidistia rather than

with typical Stegocephali. In reference to the dentition of Loxomma

Watson states that:

“The Palatine is very similar to the transverse in general character, but

bears two large tusks near its outer border. Each of these teeth has associ-

ated with it a shallow pit from which a tooth has been shed, and in which a

replacing tooth will be formed. In some cases both teeth are present: at once,

a condition which was undoubtedly only transitory; this curious type of tooth

change is very characteristic of the Stegocephalia, and is unknown elsewhere

except in the Crossopterygian ® fish, where it occurs in a very typical form in

the vomerine tusks of Megalichthys, and no doubt in many other genera, and

in Lepidosteus. This occurrence seems to me a strong additional reason for

regarding the Tetrapoda as derived from this group of fish.”

The infolded base of the teeth in the Rhizodontide is also strongly

suggestive of stegocephalian affinities, but may be only an independent

adaptive device for fastening the teeth to their bases.

6 “Throughout this paper ‘Crossopterygian’ is used as including only the three families

Holoptychiide, Rhizodentide and Osteolepide of S. Woodward's sub-order Rhipidistia,

and excluding Tarrasius, Cvelacanthus and Polypterus.” (Watson).

234. ANNALS NEW YORK ACADEMY OF SCIENCES

Although the chondrocranium of Rhipidistia, except for the basioc-

cipital and basisphenoid elements, is not well known, it was probably

fundamentally similar to that of Polypterus, and here it may be remarked

also that the sphenethmoid of that fish is similar to the element of the

same name in modern Amphibia, and that Broom (1913, p. 587) has

recently described the sphenethmoid of the stegocephalian genus Hryops

as recalling that of Polypterus in certain details.

The lower jaw of Rhipidistia differs considerably in form from that

of Stegocephali and retains a full series of gular elements median, paired

and lateral, as well as a row of infradentaries; certain of these elements,

especially the paired gulars, seem to have been lost in the Stegocephah,

while some others are with difficulty traceable.

In spite of these differences, however, the primitive stegocephalian jaw,

as described for instance by Williston (1915), Broom (1915, p. 575) and

Watson (1912, p. 11), has many important characters in common with

the jaws of Osteolepis and Rhizodopsis, as described respectively by .

Pander, Traquair and others. Putting together the suggestions of ho-

mologies made by Smith Woodward, Watson and Broom (1913, pp. 77-

78) we would have the following table:

RHIPIDISTIA STEGOCEPHALI

First infradentary Splenial (Woodward)

Intermediate infradentaries Positsplenial (Watson)

Posterior two infradentaries Angular and surangular

Dentary Dentary

Coronoids (7? “splenial’ of Amia Coronoid and precoronoid

and Polypterus)

Prearticular (Sauripterus) Prearticular

Articular Articular

STEGOCEPHALI

When the classification of the dermal elements of the skull-roof which

has been worked out in the preceding pages for the Rhipidistia is applied

to the skull-roof of the Stegocephali the following correspondence results

(cf. Fig. 3):

GREGORY, PRESENT -STATUS OF ORIGIN OF TETRAPODA 33

RHIPIDISTIA STEGOCEPH ALI

A. Dorsal series

1. Rostro-frontal segment

Premaxillze Premaxillze

Ethmoid More or less undivided Ethmoid Sharply separated

Nasals Nasals (

“Anterior frontals’” (ectethmoid) ? Septomaxillary

Frontals Frontals

2. Parietal segment

Parietals Parietals

“Squamosals” (pterotics) Pterotics (“Supratemporals”’, “supra-

squamosals’’ )

3. Occipito-nuchal segment

(Nuchal median “supratemyporal’ ) Paired dermo-supraoccipitals (post-

parietals)

Lateral “supratemporals” Tabularia (“‘epiotics’’ )

B. Cireumorbital series

Including a variable number of ele- Including always five elements: pre-

ments: in Holoptychiide, Glyptopo- frontal lacrymal, jugal, postorbital,

mide, three (‘“prefrontals’, “post- postfrontal

frontals’, “suborbitals’”) ; in Osteo-

lepidz four or five; in Rhizcdontide

five

C. Temporal-opercular series

Pterotic (‘“squamosal’’ ) Pterotic. (Ssupratemporal, suprasqua-

mosal)

Cheek plate X (—posterior postor- Squamosal

bital of Amia) and true operculum

Cheek plate X’ (interoperculum ) Quadratojugal

Operculum ? Become membranous, or transformed

into tympanic membrane of Stere-

ospondyli

Subopercuium ? Lost (become membranous)

The dorsal segments as above defined include only transverse zones ;

but, running longitudinally from the top of the orbit to the top of the

ccciput and external to the primitive spiracular cleft (Polypterus, Holo-

ptychius), there is a postorbito-pterotic row which corresponds in posi-

tion to the postorbitals, intertemporals and pterotics (supratemporals)

of Stegocephali.

336 ANNALS NEW YORK ACADEMY OF SCIENCES

The supposed correspondence in the elements of the lower jaw between

the Osteolepidx and Rhizodontidz on the one hand and the most primi-

tive Stegocephali on the other have been noted above (p. 334).

According to Watson, the resemblances in the under side of the skull

between Megalichthys and the primitive Stegocephali (Loromma, etc.),

which have a single occipital condyle, is remarkably close, and apparently

there is little doubt as to the homology of the following elements in the

two groups: premaxille, maxille, prevomers, palatopterygoids, para-

sphenoid, quadrates, hyomandibular (stapes), basisphenoid, basioccipital

Fic. 3.—Skull patterns of Trimerorhachis and Diplopterus

A, Permo-Carboniferous stegocephalian, Trimerorhachis medius (order Temnospondyli),

after Broom. B, Devonian osteolepid rhipidistian Diplopterus, after Jaekel (lettering

somewhat altered).

Abbreviations as in Fig. 2, p. 330; also: Jt, intertemporal; H.m, hyomandibular (colu-

mella auris) ; ?sy, ?symplectic (? extra columella).

In B the interfrontal foramen is identified by authors as the pineal opening. ‘The

brain was probably located far forward.

(a modified centrum). Again the elements of the chondrocranium, so

far as they are known in Stegocephali (Trimerorhachis, EHryops), seem

to correspond in general plan with the conditions in the existing

Polypterus.

It is conceivable that the Stegocephali may have branched off from

the stem of the Rhipidistia before the skull-roof became fully ossified,

but the existence of such an extensive series of correspondences offers

strong evidence of community of origin. It seems reasonable, therefore,

to conclude that the primitive stegocephalian skull has been derived from

the rhipidistian skull through the following advances:

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 337

1). The rostro-frontal segment has usually become more elongate and

always more completely differentiated (nostrils larger, separate nasals

and premaxille, distinct internal nares).

2). The orbits have increased in size and are generally displaced back-

ward farther from the nares.

3). A pineal foramen. between the parietals has appeared.

4). The elements of the skull-roof usually have acquired more polyg-

onal, or angulate, and less rectangular, outlines.

5). The postorbito-pterotic series, lying above the spiracle and bear-

ing a supraorbital sensory tract has given rise to the postorbital, inter-

temporal and true supratemporal (pterotic) of the Stegocephalia (cf.,

Stegops dwaricata Moodie).

6). The circumorbital series have come into closer sutural relations

with the surrounding elements and appear as the prefrontal, lacrymal,

jugal, postorbital and postfrontal.

7). When the shoulder-girdle became freed from the skull by the

atrophy of the posttemporals the median nuchal plate became firmly

attached to the parietals and was divided by a median suture into the

paired dermosupraoccipitals, while the lateral nuchals, also becoming

attached to the parietals and pterotics, gave rise to the tabularia (epi-

otics).

8). The operculum and suboperculum, as well as the median paired

and lateral gulars, lost their bony constituents and became membranous.

This may have resulted from a complexity of changed conditions follow-

ing upon the assumption of air-breathing habits, the dwindling of the

branchial arches and the reduction of the cleithrum (with which in

Crossopterygii the operculum is in close contact, see page 354). The

branchial chamber may also have served more or less as a resonating

chamber and sound vibrations may have been transmitted from the outer

air, through the operculum and hyomandibular (stapes)-to the side of

the otic capsule, so that the operculum may have given rise to the tym-

panic membrane of Stereospondyli, which had the locus of the operculum

and was in contact with the squamosal (preoperculum), stapes (hyo-

mandibular) and tabulare (lateral shoulder plate).

9). The preoperculum and interoperculum (X”’) covering the side and

lower part of the quadrate in Rhipidistia did not share the fate of the

operculum but gave rise to the squamosal and quadratojugal, respectively.

10). As a result of the loss of the bony character of the opercula the

fossilized skull of the Stegocephali is sharply truncated behind the occiput

and squamosal and in the young stages the branchial arches (cf., Branch-

iosaurus) are exposed. As the Stegocephali were almost certainly de-

ANNALS NEW YORK ACADEMY OF SCIENCES

rived from Osteichthyes of some sort, 1t is highly probable that their

ancestors had fully developed nuchal, opercular and branchiostegal or

gular elements, and their disappearance may be accounted for by the

foregoing hypothesis.

11). The “otic notch,” between the cornu of the tabulare and the

squamosal, was probably not a direct inheritance from rhipidistian con-

ditions but was progressively developed in the Stegocephali.

12). From Watson’s observations on Megalichthys it would appear that

the base of the skull was transmitted from the rhipidistian stem to the

earliest Stegocephali with very little change, but in the typical Stego-

cephali the median occipital condyle had been largely withdrawn and the

paired exoccipitals furnished the chief articular surfaces for the vertebral

column, while the interpterygoid vacuity, becoming much expanded, gave

rise to the well-known fenesirate palate with widely divergent pterygoids.

COMPARATIVE StTuDY OF THE PectTorAL Liwes In Priwitive FIsHEes

AND TETRAPODA

ORIGIN AND EARLY HISTORY OF THE LOCOMOTIVE APPARATUS OF

VERTEBRATES

The problem of the origin of the Tetrapoda im practice is not easy to

circumscribe or isolate: for it is almost inextricably connected with other

phylogenetic and morphological problems, cluding some of the widest

scope.

After we have compared the skulls of the earliest Tetrapods with those

of fishes and have adopted provisional views as to the homologies and

transformations of the various elements of the skull im the two elasses,

we must take up the difficult problem of the origin of paired limbs of

cheiropterygial type from some form of piscine appendages. But no

satisfactory solution of this can be attaimed until the problem of the

origin of fins in general, including both median and paired fins, has been

attentively considered. Here we must weigh Gegenbaur’s famous theory,

that the median and the paired fins have had a different mode of origin,

the paired fins being modified gill structures, against the opposing theory

that both median and paired fins have had a similar mode of origin, from

folds of skin; and after we realize the far-fetched and mystifying char-

acter of Gegenbaur’s theory and the strength of the embryological and

paleontological evidence in favor of the opposite theory that has been set

forth by Wiedersheim (1892) and in more recent years by Goodrich,

Dean, R. C. Osburn (1906) and others, we come to the further realiza-

tion that paired limbs, paired fins and median fins are all purely acces-

—

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA — 339

sory locomotive structures, which transmit to the surrounding or sup-

porting medium the thrusts of the primary and essential locomotive

organs, which are the myomeres.

If in turn we inquire into the nature and origin of the myomeres we

enter upon some of the master problems of vertebrate morphology, espe-

cially the origin of the mesenchyme, mesoderm and ccelom, the early

metamerism of the chordates and the phyletic relations of the Chordata

to other phyla of Metazoa. While no one would claim that these greater

problems are fully settled, yet the modern studies of Sedgwick, Lankes-

ter, Goodrich, Patten, Castle, Kingsley, Willey, Delage and Hérouard

(1898) and many others, afford a strong evidential basis for some such

synthetic concept of the earlier evolution of the chordates as may now be

outhned.’

The pre-Silurian and perhaps pre-Cambrian ancestors of the Chordata

were, I believe, related neither to the Arthropoda, Annelida or any other

phylum exhibiting metamerism; the elaborate resemblances discerned by

Patten and others between limuloid and chordate structures being re-

garded as the homoplastic results of similar locomotive adaptations on

the part of independent phyla having in common chiefly the following

characters: (a) anteroposterior motion, (0) a metameric repetition of

mesodermal tissue, (c) a subsequent independent process of cephalogen-

esis or concentration of neuromeres.

Possibly these pre-Silurian chordates may have traced back their origin

to the stem of the echinoderms, or they may have been ccelenterates of

some sort, as suggested by Sedgwick and by Masterman. Possibly they

are pictured in a general way by the early larve of Balanoglossus and

echinoderms. At any rate they were at first more radiate than bilateral

in plan. They had a more or less ciliated epidermis, the cilia being the

first locomotive organs of all known phyla. Their primitive gut or

archenteron may have borne several diverticula, more or less similar to

the five “archimeres” of Balanoglossus, which were destined to give rise

to the myoccelomic pouches of chordates (Lankester). These myocce-

lomic pouches are thought by some to have surrounded the gonads, which

were derived from the archenteron, and a primary segmentation of the

gonads, corresponding to that of the myotomes, was assumed ; but Kings-

ley states (1912, p. 319) that in the existing vertebrates no metamerism

of the gonads exists. At any rate the segmental myoccelomic pouches

very early came into functional relations with the gonads, through the

formation of segmental nephridial ducts.

7A preliminary report on this subject was read before the New York Academy of

Sciences, Nov. 13, 1911. Abstract in Science N. S., vol. XXXIV, p. 892. 1911.

340) ANNALS NEW YORK ACADEMY OF SCIENCES

The steps by which the primitive radiate symmetry changed into bilat-

eral symmetry are quite vague, but the change no doubt involved the

progressive contractility of the myoccelomic pouches and the assumption

by some of them of a partly locomotive function.

Bilateral symmetry was, at least, one of the earliest of all chordate ac-

quirements and may have led the way for such fundamental characters

as metamerism, a notochord and a nervous system of vertebrate type.

The myoccelomic pouches very early began to increase in size and num-

ber and in power of contractility; all this accompanying an emphasis of

bilaterality, a moderate lengthening of the anteroposterior axis and the

assumption of an obtusely fusiform shape. The nervous elements, per-

haps originally more or less diffused in the skin, were segregated in defi-

nite tracts, which foreshadowed the chordate neuron and segmental

nerves. The anterior part of the gut gave off paired diverticula on either

side, which may at first have served to draw in a food-bearing current of

water, but later assumed a respiratory function, acquired exterior fistule

and gave rise to the branchial apparatus of chordates. ‘The myoce-

lomic sacks, extending ventrally, inclosed the primitive gut below; the

ccelenterate mouth was closed and a new mouth was opened, formed from

the coalescence of opposite pharyngeal diverticula. The dorsal moiety of

the myoccelomic sacks gave rise to the muscle segments, the ventral moi-

ety to the ccelom. Meanwhile the mesenchyme, which in modern verte-

brates arises chiefly from the splanchnic and somatic walls of the muscle

plates, was giving rise not only to the corpuscles of the blood and lymph,

but also to the deeper layer of the skin (corium), the involuntary muscles

and the connective tissue antecedents of cartilage and bone (Kingsley),

all of which were destined to become of the utmost importance in the

further development of the locomotive organs.

The notochord perhaps arose as a ciliated groove on the dorsal wall of

the gut, its locomotive function being secondary. The circulatory, excre-

tory and respiratory structures were all accessory adaptations for more

rapid metabolism, following the primary change of myoccelomic pouches

into locomotive organs. During these early stages there was but little

differentiation of head and trunk, the pharyngeal region was large, and

cephalogenesis, or fusion of neuromeres etc., was in a low stage.

A critical stage in vertebrate evolution was reached when, through the

codperation of the vascular system and of the rapidly differentiating

mesenchyme, a twofold skin was substituted for the primitive ectoderm,

and connective tissue began to be formed around the notochord, around

the sensory capsules, beneath the skin, between the myomeres and be-

tween the gill pouches. Still later the process of ossification and calezfi-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 341

cation not only gave rise to a more fully protective exoskeleton, but af-

forded more rigid bases or fulcra for the attachment of the segmental

muscles.

A further development.of the muscular system, involving superior

locomotive powers, rapidly ensued, resulting in a corresponding improve-

ment in offensive and defensive adaptations. The primitive food habits

required only that microscopic food particles be drawn into the pharynx

by ciliated tracts and later by the sucking action of the branchiai

pouches, but these primitive habits were later abandoned for the active

pursuit of larger prey. At this stage also the conditions necessary for

fossilization were fulfilled, the paleontological record opens with the Si-

Iurian ostracoderms, and speculation can be replaced by recorded history.

OSTRACODERMI

The subsequent history of the head, as thus conceived, has been out-

lined above (pp. 318-338). The acquirement of a many-layered skin and

of osseous or calcareous deposits in the exoskeleton also conditioned the

formation and rapid improvement of accessory locomotive organs, espe-

cially the fins. From the beginnings of bilateral symmetry and of the

differentiation of the head-end from the tail-end, there must have been a

tendency for the locomotive end of the body to protrude behind the ter-

minus of the primitive gut, and to become laterally compressed, while the

tip of the head-end became either dome-shaped or depressed In the

ostracoderms the caudal end is already provided with a web of skin, serv-

ing as a sweep and strengthened dorsally by ridge scales. The form of the

hard parts was everywhere conditioned by the arrangement of the under-

lving myomeres and their connective tissue septa, as shown clearly in the

arrangement of the scales in modern fish. Dorsal, ventral and caudal

outgrowths of various shapes, strengthened by hard scales or scutes,

served to transmit the thrust of the myomeres to the surrounding medium.

The fusiform free-swimming Birkeniide while well provided with

median or vertical fins were apparently not provided with horizontal or

paired fins for steering up and down, this perhaps being effected by twist-

ing the body. Thelodus and Lanarkia on the contrary were more or less

ray-like in form and the lateral angle or lappet of the body may have

assisted in steering up and down and in veering or dipping to one side or

the other. The Cephalaspide had a pair of fleshy, scaly flaps behind the |

cephalothoracic shield, which may have been of considerable assistance

in steering. In the Antiarchi the pair of lateral appendages behind the

head were protected by a many-layered osseous armor, jointed so as to

permit bending, and vaguely suggestive both of arthropod appendages

349 ANNALS NEW YORK ACADEMY OF SCIENCES

and of the shell-covered flippers of tortoises. A few authors have even

endeavored to homologize the paired appendages of Antiarchi with the

pectoral limbs of gnathostomes! Pelvic paired fins are absent in the

Ostracodermi and Antiarchi, and also in the Arthrodira, as Dean eae

pp. 282-287) has shown.

ARTHRODIRA

The exoskeleton of the Antiarchi afforded an ample base for their

paired appendages as well as for the muscles of the head and thorax, and

the same is true in the case of those Arthrodira which, like Acanthaspis,

had paired spine-like appendages attached to the anteroex.ernal corners

of their osseous plastron. Here it may be noted that the attempts of

Jaekel, Tate Regan and Patten to homologize the elements of the thoracic

plates of Arthrodira with the dermal plates of the shoulder-girdle of

Osteichthyes appear to the present writer to constitute a begging’ of the

question. There are, it is true, vague resemblances between the “clavicu-

lar” and other thoracic plates of Arthrodira and the pectoral plates of

Osteichthves, but in view of the amazing powers of convergent evolution,

which are known by experience to many investigators, why should we

assume a homology or a series of homologies and then regard them as

a basis for phylogenetic speculation? The “homologies” assumed by

Jaekel, Patten and Tate Regan are based merely upon a general similarity

in the spatial relations of certain plates in Arthrodira and Osteichthyes

with reference to assumedly homologous starting-points. Given paired

orbits in the two phyla and Jaekel will call the median dorsal element

lving between the orbits “frontale,’ the elements behind it “parietalia”

and the median occipital element “‘occipitale superius’ and assume that

the so-named plates are homologous with those of Osteichthyes. And the

supposed homologies of the arthrodiran shoulder plates with those of

Osteichthyes are equally arbitrary and unconvincing. This matter is of

some importance to our main topic, the origin of the Tetrapoda, for in

Professor Jaekel’s early schemes the Tetrapoda and the Placodermi were

indicated as having sprung from a common pro-tetrapod stock; Patten

too connects the Arthrodira with the stem of the Dipnoi and Amphibia

and homologizes the pectoral plates of Dipnoi with those of Arthrodira.

According to the view here adopted the Arthrodira may be an offshoot

_ from the antiarchian stem, which “‘paralleled” and even surpassed the

Osteichthves in the development of the exoskeleton of the head and

thorax, but failed to build up the endoskeleton to the same degree. Nor

were their accessory locomotive organs (caudal fin, dorsal fin) as highly

crganized as in either the Elasmobranchii or the Osteichthyes.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 343

ELASMOBRANCHII

In the earliest known elasmobranchs, the Acanthodii of the Upper

Silurian and Devonian, the locomotive apparatus as a whole was in a

much more advanced stage than in the typical Ostracodermi or Arthro-

dira. The locomotive part of the body, namely, the trunk and tail, is

from three to six times as long as the cephalo-pharyngeal region and no

heavy thoracic armor impedes the undulation of the fusiform body. In

the earliest forms the median and paired fins are of large size and

unusually numerous, for in addition to the two dorsal fins, the anal and

the caudal fin, there is a whole row of accessory paired fins, or fin spines,

between the pectorals or pelvics, suggesting the former presence in this

region of paired ventral fin-folds. The Acanthodii are, however, defi-

nitely excluded from ancestry to the higher types, by the fact that the

exoskeleton was more highly developed than the endoskeleton. The an-

terior borders of all the fins both median and paired were supported by

fin-spines, which are believed (Dean, 1907, p. 216) to represent clusters

of originally metameric dermal tubercles. Functional pectoral and pelvic

eirdles were also developed from dermal elements, and perhaps served for

the insertion of powerful muscles, as well as for the support of the heavy

spines. But the underlying cartilaginous elements were little if any

developed and in my view the dermal elements of the girdles were

analogous but not homologous with those of Osteichthyes. In some

acanthodians (Gyracanthus) the pectoral fin-spines became overspecial-

ized and attained a relatively enormous size; in other lines all the spines

were reduced and the body in the later tvpes became much elongate as in

many other decadent groups of fishes (A. 8. Woodward).

In brief the Acanthodiu failed to carry the exoskeleton beyond a low

stage of evolution, and their dermal shoulder-girdle was developed inde-

pendently of that of the Osteichthyes.

The Cladoselachii avoided the line of specialization typified by the

Acanthodii and indeed went to the other extreme in sacrificing a large

part of the exoskeleton. But they carried much further a process which

in the Acanthodii was barely begun and soon abandoned, namely, the

building up and calcification of rods of cartilage, lying between the

myomeres and extending out into the median and paired fins; these

metameric cartilaginous rods reached almost to the tips of the fins, they

were jointed at the body line and within the trunk they underwent more

or less coalescence and enlargement, giving rise to the cartilaginous

girdles, basals and radials. In this group the paired fins, like the median

fins, had a wide base which was not exserted posteriorly from the body :

344 ANNALS NEW YORK ACADEMY OF SCIENCES

nor were the basal cartilages widely protruded. Consequently the paired

fins were used rather as steering planes, which could be raised or lowered

or gently undulated, rather than as paddles, capable of twisting about a

narrow base.

The typical elasmobranchs have followed a conservative line in the

evolution of their fins: in the distal portions of all the fins they have

developed horny dermal rays or ceratotrichia, which lack a bony or cal-

careous basal portion, but serve well as the flexible elastic fin-web. In

the typical sharks the basal cartilages have become more or less widely

protruded from the body-wall, the base has shortened and the posterior

border of the fin has become sharply exserted or entirely freed from the

body-wall, with more or less rearrangement of the basals and radials;

consequently the pectoral fins of sharks and still more of chimeroids have

become very efficient paddles, capable of a wide range of movements. In

the skates, on the other hand, after the primary shortening of the base

and coalescence of some of the basal rods, there was a secondary antero-

posterior widening of the fin, a multiplication of the radials, with dichot-

omization of the distal ends, and a great forward and backward exten-

sion, with corresponding emphasis of the power to undulate the outer

border. The pectoral girdle accordingly becomes a stout depressed hoop

and secures a firm articulation dorsally with the vertebral column. All

such improvements in the median and paired fins have been accompanied

by a great strengthening of the axial skeleton, and by the functional

replacement of the notochord by the calcified centra, developed in and

around the perichordal sheaths.

The pleuracanth sharks are of historical importance in any discussion

of the early history of the paired limbs. Their pectoral limbs, recalling

the “archipterygial” type of Gegenbaur, were assumed to be the most

[Timitive form known and the vague resemblance of the pectoral girdle

and fin to a gill-arch and its extrabranchial rays were by no means over-

looked. But it is now coming to be realized that the pleuracanths, which

are of Permocarboniferous age, were highly specialized, aberrant sharks,

living in fresh-water, along with branchosaurs, microsaurs and other

swamp-dwelling types, wriggling about with their long Gymmnetus-like

body or paddling with their Ceratodus-like pectorals. Whether the

diphycercal tail of pleuracanths is a primitive structure is doubtful.

The loss of the primitive heterocercal tail and the assumption of the

gephyrocercal form in Ceratodus, Protopterus, Lepidosiren and certain

swamp-living teleosts (symbranchoids, gymnotids, gymnarchids, etc.) is

an indication not of primitiveness, but, as Dollo has shown, of degen-

erative specialization. On the other hand the tail-fin of pleuracanths

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 345

approaches the hypothetical generalized form which Schmalhausen (1913,

¢. 66), after thorough studies of the musculature and skeleton of the

caudal fin of fishes, regards as antecedent to the heterocercal tail of

typical elasmobranchs.

That the pleuracanths are truly sharks, although of a peculiar order,

is definitely proved by their skull-structure, which has been cleared up

by Hussakof (1911). As for their pectoral fins, the general resemblance

to those of Ceratodus offers an excellent example of convergent evolu-

tion: the protrusion of the metapterygial axis, perhaps followed by a

process of asymmetrical budding, having resulted in both phyla in a

more or less mesorhachic or biserial fin; so that we may regard this type

as a “morphon” which has arisen independently in widely removed phyla.

The pelvic fins of Pleuracanthus, and indeed of all sharks, are, according

to the view here adopted, in a lower stage of evolution, since their bases

are less widely protruded from the body and they have departed less

from the fin-fold type.

The large and high coracoscapular cartilage of pleuracanths remains

separate from its fellow of the opposite side; in form it parallels that of

Tetrapoda and supports the large pectoral limbs, which have a single

proximal basal piece analogous to the humerus. If the pleuracanths had

happened to develop dermal plates around the shoulder-girdle we should

inave had still more resemblances to Ceratodus and the Tetrapoda, to lead

further astray investigators who neglect the “potency of convergence.”

In brief the elasmobranchs show a marked advance over the ostraco-

derms in all locomotive adaptations. In the development of median and ~

paired fins the earliest elasmobranchs overemphasized the exoskeleton ;

some of the later ones on the contrary neglected the exoskeleton (Clado-

selachii, pleuracanths), but developed the endoskeleton to a high stage.

The basal cartilages, which were formed between the myomeres of the

fins, coalesce, enlarge and are widely protruded from the body to form

vectoral paddles either of tribasal, or rarely, of sub-mesorhachic type.

The pelvics remain on a lower stage of evolution, retaining usually a

broad base and acquiring an over-extended metapterygium.

ACTINOPTERYGII

The earliest Actinopterygii preserved that fortunate balance between

endoskeletal and exoskeletal structures which, as Dr. Smith Woodward

(1906) has shown, was essential for the highest development. First they

either inherited or reinvented bone-cells, which, being carried to almost

any desirable point by the mesenchyme and vascular system, served to

reinforce the exoskeleton and to replace the cartilaginous endoskeleton.

346 ANNALS NEW YORK ACADEMY OF SCIENCES

The Actinopterygii also introduced several other important improve-

ments in the accessory locomotive structures. In the exoskeleton they

avoided overspecialization of the outermost layers, a cul-de-sac into which

the elasmobranchs entered, and while at first developing in due propor-

tion the ganoine, cosmine, vasodentine and isopedine strata, they avoided

the errors of the typical ostracoderms and antiarchians and did not

cumber themselves with a massive carapace and plastron.

In the median and paired fins they were fortunate in evolving a type

of rod-like scales, which by fusing end to end, gave rise to the dermal

rays; these soon became the most important part of all the fin-web and

entirely superseded the horny fin-rays of an earlier period. With the

advent of these superior fin-rays and with the concomitant strengthening

of the vertebral column by neural and hemal rods, and finally by ossified

centra, the caudal fin in the Actinopterygii became of predominant func-

tional importance and changed from the heterocercal to the homocereal

type, receiving the powerful thrusts of the myomeres, which were trans-

mitted to it along the reinforced vertebral column. The hemal rods

below the caudal column served to link the tail to the backbone. At first

they were slender and numerous, but gradually were reduced in number

and expanded into the broad hypural bones, around which the stout

dermal rays were tightly clamped.

In the later Actinopterygii the body frequently became elongate, the

paired dorsal and anal fins were reduced, the dermal rays became reduced,

and the once powerful caudal degenerated into the pointed gephyrocercal

type. The varied history of scales and dermal rays of the Actinopterygii

illustrates the comparative rapidity with which these structures change

or disappear. The forerunners of the group may well have had scales

like those of the most primitive crossopterygians and dipnoans; but even

in the Paleoniscide the ganoine has become many-layered and the cos-

mine layer is modified. In later Actinopterygii the ganoine and cosmine

disappear, the scales sink beneath the skin, lose their osseous tissue and

become horny.

The over-development of the lepidotrichs or dermal rays may perhaps

be responsible for the failure of the Actinopterygii to attain the highest

development of the endoskeleton of the median and paired fins. In the

earliest form (Cheirolepis) as restored by Smith Woodward the pelvic

fin had a very extended base with short rod-like basals and radials, the

pectoral fin had a shorter base, which was, however, wider than that of

later types; the dermal rays were long, numerous and scale-lke. With

the collapse of the archipterygial theory there is no longer any reason

why this wide-based type of fin that occurs so near the beginning of the

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 347

record, shou!d be regarded as a highly modified archipterygium, especially

as the paired fins are identical in plan with the vertical fins. And yet

some authors (e. g., Braus, 1901) have assumed that the paired fins of

Amia, a much later and more specialized actinopterygian than Chetro-

lepis, were of reduced archipterygial type. The point is important in

relation to the origin of the Tetrapoda, for if the early Actinopterygu

ever passed through an archipterygial stage they might be more closely

related to the Tetrapoda than is generally admitted. But none of the

Fic. 4.—Pectoral girdle of Acipenser and Amia

(A) Acipenser sturio, after Goodrich, from Gegenbaur. (B) Amia calva, from Good-

rich. Inner (medial) view of right side.

Dermal elements: cv, clavicle (“‘infraclavicle”’) ; cl, cleithrum (‘clavicle’) ; s.cl, supra-

cleithrum ; p.t, posttemporal; po.cl, postcleithrum.

Cartilage elements: «0, coracoid; m.c, mesocoracoid; gl, articular surface: s¢, scapula ;

d.c, dorsal cartilage.

The sturgeons have the largest coracoscapula of any Actinopterygii: it is unossified ;

the true clavicles are present and of large size. In Amia, as in more typical Actinop-

terygii, the coracoscapula mass is of relatively small size, this corresponding with the

relatively small size of the fin-muscles; the coracoscapula is more or less ossified and

suturally divided into dorsal and ventral moieties named scapula and coracoid, respect-

ively ; the true clavicles have disappeared, and the cleithrum is connected with the skull

by the stout supracleithrum and forked posttemporal.

early Actinopterygii show the least visible trace of ever having had widely

protruded basal elements in the paired or median fins: on the contrary

they emphasized the dermal rays and an undulatory movement of the

margins of the fin, rather than of its fleshy base.

In the shoulder-girdle (Fig. 4) the Actinopterygii also developed the

dermal elements rather than the cartilaginous endoskeleton, the sturgeons

and their allies being the only ones with a large coracoscapula cartilage

(Fig. 4A). In most Actinopterygii the coracoscapula cartilage is small

348 ANNALS NEW YORK ACADEMY OF SCIENCES

(Fig. 4B), while the dermal elements of the shoulder-girdle (except the

true clavicles which disappear) are fully developed. ‘These dermal ele-

ments are associated with the opercular and branchiostegal series and

serve to attach the pectoral girdle to the skull; but their chief function

is to serve as a base for the coracoscapula, which bears the principal

muscles of the fin. The homology of all these elements with those of

the Tetrapoda is discussed below.

In brief the typical Actinopterygii while progressively emphasizing the

bony tissue, both in the exoskeleton and the endoskeleton, enjoyed only a

moderate development of the basals and radials of the paired fins, never

protruded the basal elements widely from the body-wall and depended

primitively on the caudal fin as the chief accessory locomotive structure,

originally using the other fins chiefly as keels, rudders, brakes and bal-

ancers (R. C. Osburn, 1906).

DIPNOI

In this group the exoskeleton is primitively like that of the earliest

Crossopterygu, but it soon undergoes degenerative changes (Goodrich,

1909, pp. 230, 238) sinking beneath the skin and losing the ganoine and

cosmine layers. The endoskeleton, on the other hand, never passes

beyond the cartilaginous stage.

The earliest Dipnoi resemble their contemporaries the Rhipidistia in

having two dorsal fins, a heterocercal tail, lepidotrichia with ganoine and

cosmine, and paired fins of mesorhachic or biserial type.

The tail never attains the homocercal type; it never attains much-

expanded hypurals, but passes from the primitive heterocercal type

through intermediate stages, described by Dollo (1895), into the degen-

erate gephyrocercal. In such an animal the caudal fin has not the com-

manding functional importance that it has in the typical Actinopterygu,

and there is a very evident tendency to throw the function of locomotion

more upon the paired fins and upon an eel-like undulation of the body.

The Dipnoi parallel the recent Amphibia in this respect and the more

specialized types also show a reduction of the dermal rays. The paired

fins of the Devonian Dipnoi and still more of the modern Ceratodus are

externally very unlike the paired ‘fins of elasmobranchs or Actinopterygu;

they also resemble vaguely the paddles of plesiosaurs rather than the

cheiroptergia of tetrapods. The modern Protopterus, however, some-

times uses its pectoral and pelvic fins as if they were legs, crawling about,

while floating in the water, on the tips of them (Dean, 1903) : while

Ceratodus sometimes rests in the water (Dean, 1906), with the tips of

the pectorals turned downward and touching the bottom. But the ability

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 349

to do this ranks very far below the ability to support the weight of the

body on the paired limbs without the buoyant effect of the water, as in

the Tetrapoda.

In Ceratodus the preaxial or upper border of the pectoral fins is be-

heved to be serially homologous with the ventral border of the pelvics.

Braus (1901, p. 165) states that the earliest anlage of both fins are

horizontal, that the nerve entrance of the pectoral is on the lower, or

ventral, surface, while that of the pelvic is on the dorsal surface. Ac-

cording to this view the preaxial border of the pectoral has been rotated

upward as in most Actinopterygii while in the pelvics the preaxial border

has been rotated downward. Schneider (1886, quoted by Howes, 1887,

p. 12) held that “die Seitenstrahlen der dorsalen und ventralen Halfte

der [pectoral and pelvic] Flossen sind ungleich,” and that the “Seiten-

strahlen der dorsalen Halfte der emen Flosse entsprechen derjenigen der

ventralen Halfte der anderen.” Goodrich (1909, p. 244) says that “when

at rest the preaxial margin of the pectoral fin is borne upward; the re-

verse is the case with the pelvic fin.” But Howes (1887, pl. II) has de-

scribed a pelvic fin of Ceratodus in which this reversal has not been

effected and he records so many irregularities in the structure of both

pectoral and pelvic fins that the condition of reversed homology seems to

be incompletely attained.

Many investigators (cf. Keith, 1912, p. 418) have sought to show that

in man there is a somewhat similar condition of reversed homology in

the borders of the pectoral and pelvic extremities; but according to H. H.

Wilder (1909, p. 245), the evidence of embryological history as well as

ef comparative anatomy lends strong support to the opposite view that

in the Tetrapoda the preaxial or anterior border of the pectoral extremity

is homologous with the preaxial border of the pelvic extremity, a con-

clusion which is further strengthened by the general correspondence in

the arrangements of the elements of the manus and pes in the most primi-

tive Tetrapoda of the Permocarboniferous, such as Hryops, Stereosternum

and the cotylosaurs.

If the latter view be correct the reversed homology of the borders of

the pectoral and pelvic fins in Ceratodus is a point of marked difference

from the Tetrapoda, which may have been acquired only by the later

Dipnoi. |

As the paired fins of even the oldest Dipnoi are already biserial in

form, there is no direct paleontological evidence as to their origin. In

the allied group of Rhipidistia, however, we get several hints as to the

origin of the “archiptervgial” type. Osteolepis, which I regard as the

most primitive member of the group, on account of its skull structure,

250 ANNALS NEW YORK ACADEMY OF SCIENCES

rhombic scales and heterocercal tail, has a short wide fleshy lobe in the

pectorals while the pelvics according to Traquair’s restoration (Good-

rich, 1909, p. 283) have still shorter and relatively wider lobes. Thus

the pelvics in turn are only a little more advanced than the anal and

posterior dorsal. In Glyptopomus the pelvic base is narrower and the

axis more protruded, but the resemblance to the widely based dorsal and

anal fins is still obvious. But the pectoral of this genus has become

almost fully archipterygial. Coming to the Dipnoi, in Dzpterus the

process is carried further and the resemblance between the pelvic and the

anal is progressively effaced. In the latter Dipnoi the “archipterygium”

is perfected by the protrusion of the mesopterygial axis to the extreme

tip of the fins; the pelvic fin also is finally as fully developed as the

pectoral.

Turning to the evidence of comparative anatomy we find that in

Ceratodus the plexuses of nerves that supply the pectoral and pelvic fins

are brought together by the coalescence of many segmental nerves (Braus,

1901) and by analogy with the conditions in sharks (cf. Goodrich, 1909,

pp. 72, 78), where both the paired and median fins are formed in the

same manner, through the concrescence of myotomic elements, it appears

highly probable that in Ceratodus also the narrow-based fins arose through

the concentration of metameric elements, the base becoming constricted,

as the power of rotating the fin increased. From the cone-in-cone ar-

rangement of the myomeres of the Ceratodus paired fins it seems also

probable that the outgrowth of the mesopterygial axis to the very tip of

the fin was due to a sort of apical budding, or repetition of similar seg-

ments, on the part of the mesopterygium and its radials. That the cen-

tral axis of the Ceratodus fins does represent an outgrowth of the mesop-

terygium was held by Huxley in opposition to Gegenbaur, and was

supported, with strong evidence, by Howes (1887).

From these and similar considerations I reject the traditional view

that the “archipterygia” of Dipnoi are primitive structures and I regard

the imperfect archipterygia of the Devonian Osteolepis as more primitive

than the perfected archypterygia of the modern Ceratodus. Nor can I

accept the views of Watson, Smith Woodward and others that the paired

limbs of the Rhipidistia are “reduced archipterygia”; they seem rather

to be incompletely evolved or primitive “archipterygia,’ with less ex-

tended mesopterygia and an asymmetrical arrangement of the radials

(parameres), those on the postaxial border being fully developed, the

preaxial parameres being small or wanting.

The pectoral and pelvic fins of existing dipnoans iatee this important

character in common with the paired limbs of Tetrapoda, namely, that

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 351

the endoskeletal base consists of a single proximal element, analogous

respectively with the humerus and femur. Other proposed homologies

of the more distal elements with the forearm, carpus and digits have been

assumed, but lack adequate evidential support.

The shoulder-girdle of the Devonian Dipnoi evidently included dermal

elements (“infraclavicles,” “clavicles,” etc.) and underlying cartilaginous

elements, which, as Smith Woodward says (1898, p. 63), “seem to have

been always too slightly calcified for preservation.” But from the re-

semblance of the pectoral limb of Dipterus to that of Ceratodus, it is

highly probable that the endoskeletal elements in the Devonian forms

were fundamentally similar to that of Ceratodus.

The accompanying figures (Fig. 5), drawn from a specimen of Cera-

todus forstert by Professor L. A. Adams, show that in the modern genus

the chief cartilaginous element, which is supposed to be homologous with

the coracoscapula of the Tetrapoda, is of large size, and includes a short

ascending ramus corresponding to the scapula, a tripartite, protuberant

articular pedicle, and a long forward-and-inwardly projecting coracoid.

This undivided element is firmly adherent to the antero-internal surface

of the conjoined dermal elements first named by Gegenbaur clavicle

(“infraclavicle” of early writers) and cleithrum (“clavicle”). A small

median ventral cartilage (m. v. c.) unites the opposite clavicles, while a

broad dermal supracleithrum connects the pectoral girdle with the post-

temporal.

This shoulder-girdle is apparently more primitive than that of the

primitive Tetrapoda in the slight dorsal extension of the scapula, in the

unexpanded form of the coracoid, and especially in the protuberant form

of the articular region; also, there are no centers of ossification tending

to divide the coracoscapula mass into distinct elements. The dermal

elements, especially the cleithrum and supracleithrum, are of large size,

whereas in Tetrapoda they are much reduced. This shoulder-girdle is

on the whole nearer to the tetrapod type than that of any other existing

fish.

The pelvic girdle of Ceratodus also differs from that of typical fishes

and approaches that of the primitive Tetrapoda in having a stout carti-

lagious base, perhaps homologous with the ischio-pubis. The “pre-

pubic” and “epipubic” processes are of doubtful homology. No dorsal

process analogous with the ilia and extending up from the acetabulum

toward the backbone is present.

In brief the more specialized Dipnoi have paralleled the urodele Am-

phibia in the reduction of the dermal rays, in the degeneration of the

scales, in the non-development of a powerful homocercal tail, and in the

352 ANNALS NEW YORK ACADEMY OF SCIENCES

Pp 0. AX.

Fig. 5.—Pectoral girdle and fin of Ceratodus forsteri

1, Left half of girdle, with fin: outer side seen somewhat from below, showing car-

tilaginous (stippled) and dermal elements; 2, inner side; 3, posterointernal view, show-

ing left coracoscapula and articular surface for pectoral fin; 6, posterointernal view

after removal of coracoscapula; 7, front outer view; 4, rear view of left coracoscapula

cartilage; 5, outer side of same, with articular surface for cleithrum (cf. no. 1).

Dermal elements: cv, clavicle; cl, cleithrum ; s.cl, supracleithrum.

Cartilaginous elements: co.sc, coracoscapula; m.v.c, median ventral cartilage; gl, ar-

ticular surface for “humerus”; H, “humerus” (proximal basal).

pr.az, preaxial or dorsal border; po.az, postaxial or ventral border.

Unlike that of the Tetrapoda, the stout coracoscapula cartilage has a very short

ascending ramus; the glenoid region is widely protruded and the muscles are better

developed on the visceral or internal surface.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 353

assumption of an eel-like habitus. On the other hand the pectoral and

pelvic girdles of the more primitive Dipnoi appear to preserve certain

characters which may be truly homologous with those of the ancestral

Tetrapoda. The “archipterygia” of Dipnoi are primitive only in name,

although evolved in their essential features at an early epoch. That such

“archipterygia” are ancestral to the cheiropterygia of tetrapods is un-

proved and rather unlikely. The single proximal basal pieces of the

paired fins may possibly, however, be homologous with the humerus and

radius respectively; although it is quite conceivable that here too the

resemblances may be only homoplastic and due to similar concrescence

of several originally separate basal pieces.

CROSSOPTERYGII

The existing Polypterus has no doubt departed widely from the primi-

tive Devonian Crossopterygiul, both in the exoskeleton and in the endo-

skeleton and has paralleled the Actinopterygii in many respects. Among

its progressive characters may be noted: the replacement of the notochord

by ossified centra, the development of osseous neural and hemal arches

and hypural bones (the latter, however, being arrested in development),

the reduction and ossification of the coracoscapula and its division into

two elements (Fig. 6), the reduction of the radials and basals, the fusion

of the basals of the pelvic fins to form a pelvis, which is ossified. And

Polypterus has also become highly ichthyized in the brain and many other

characters of the soft anatomy. Among its peculiar or aberrant special-

izations are to be reckoned the gephyrocercal tail, the subdivided dorsal

and the multiplication of the radials in the pectoral fin.

The proximal elements of the pectoral fin are often compared with the

pro-, meso- and metapterygia of sharks; the so-called “pro-” and “meta-

pterygium” forming a V-shaped articulation with the scapulo-coracoid ;

the ovoid “mesopterygium” lying in the middle of the fleshy lobe of the

fin; but this arrangement may rather represent a modification of the

Eusthenopteron and Sauripterus type of fin described below. Possibly

the “metapterygium” and “propterygium” may be homologous with the

radius and ulna of Sauripterus; the fused “mesopterygium” may repre-

sent the fused mass of radials, or parameres, which in Sawripterus con-

verge toward the central axis. The single proximal piece (true meso-

pterygium of Sauripterus) in Polypterus has either been lost or has per-

haps fused with the coracoscapula. Indeed the whole coracoscapula in

the larval Polypterus (cf. Goodrich, 1909, p. 296) bears a puzzling re-

semblance to the humerus of Sauripterus.

At any rate the pectoral of Polypterus is further removed from the

354 ANNALS NEW YORK ACADEMY OF SCIENCES

primitive cheiropterygium than is the pectoral of Sauripterus. Klaatsch

(1896) would derive the cheiropterygium of Amphibia from the crossop-

terygium of Polypterus, but the lack of a single basal piece or humerus,

as well as the secondary multiplication of the distal radials, are aberrant

specializations. However the pectoral of Polypterus does suggest the

tetrapod type in this respect: that, as Klaatsch and Budgett have shown,

it can be turned downward almost like a hand, so that the larval fish

sometimes rests on the palmar surface of the dermal rays; the pectoral

Fic. 6.—Pectoral girdle of Polypterus bichir

A, Left side, outer view. B, Posteroventral view.

Dermal elements: cv, clavicle; cl, cleithrum; s.cl, supracleithrum ; po.cl, postcleithrum ;

p.t, posttemporal.

Cartilaginous elements: co. coracoid: sc. scapula.

This modern fish retains the true clavicles of the Devonian crossopterygians and

tetrapods. Its scapulocoracoid, however, parallels that of more typical fishes and affords

a short, wide base for muscles that raise and lower the fin. The forked posttemporal

and wide supracleithrum serve to connect the girdle with the skull; while the marked

concavity of the whole anterior margin of the girdle denotes the presence of a large

operculum and large branchial arches.

is also well provided with muscles for elevating, depressing, protracting,

retracting, abducting, adducting and twisting the fin (Klaatsch, Braus).

Before considering the relationship of the Rhipidistia to the Tetrapoda

it is necessary to decide which is the most primitive family, and what

has been the trend of evolution within the group. Smith Woodward

(1898, pp. 71-80), perhaps influenced partly by the view that the

“archipterygial” type of fin is primitive, seems to have regarded the

Holoptychiide as having the most primitive paired fins, and the Rhizo-

dontide as more progressive in the “abbreviation” of the paired fins.

“Through Rhizodopsis and Gyroptychius,’ he said (p. 77), “the Rhizo-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 355

dontide pass almost imperceptibly into the Osteolepide, which have

thombic scales and usually exhibit the fusion of the bones in the frontal

portion of the skull and in the mandible.” Goodrich (1909, p. 285), on

the other hand, places the Holoptychiidz and the allied Glyptopomide

first, the Osteolepide next, the Rhizodontide third and the Onycho-

dontide last. He says (p. 285): “It is evident from the structure of

the paired fins that the Rhizodonts differ considerably from the previous

three families | Holoptychiide, Glyptopomide, Osteolepide], and they

should perhaps not be included in the same order.”

The Osteolepidz present the following assemblage of characters which

in view of all that has been said above I must regard as primitive.

1). Their paired fins are obtusely lobate, 7. e., incompletely “archipterygial’’

and less different from the median fins, in contrast to the extremely

acute “archipterygial” fins of Holoptychiide.

2). The scales are rhombic, like those of the earliest Actinopterygii, and they

retain the fully developed ganoine and cosmine layers, which are pro-

gressively lost in the Rhizodontide and highly modified in the Holopty-

chiide (cf. Goodrich, 1909, pp. 217, 285).

3). In the Osteolepide the dermal rays of the median and paired fins are less

produced than in the Rbizodontide.

4). The caudal fin in Osteolepis is of a primitive heterocercal type, whereas in

Rhizodontidz it externally approaches the homocereal type (through

Gyroptychius ).

5). Osteolepis had a persistent unconstricted notochord without ossified centre,

but the larger and more progressive members of both families (Megu-

lichthys, Eusthenopteron) had ring centra.

6). As to the rostrum of Osteolepis representing a “fusion’’ of elements I have

given reasons above (p. 327) for the contrary view that the nasals,

dermethmoids and other elements had not yet been divided by sutures.

In brief Osteolemis appears to offer an ideal stem form for several

phyla. One line with progressively cycloidal scales and intermediate

characters of the skull-top seems to have led through Gyroptychius (fig-

XXXII) and Husthenopteron (Hussakof, 1912, Fig. 2). Another line,

preserving the rhombic scales, perhaps passed through Diplopterus (Pan-

der, 1860, Tab. 4) and culminated in Megalichthys (Wellburn, 1900, pl.

XIII; cf. our Fig. 7). The skull of Rhazodopsts, as figured by Traquair

(1881, Figs. 1, 2) seems to be allied in pattern to that of Megalichthys,

as figured by Wellburn (1900, pl. XV) and yet the former is referred

to the Rhizodontide, the latter to the Osteolepidze: and there are other

indications that these two families are very closely allied.

In the Holoptychiide the scales have become large, fully cycloid and

deeply overlapping and have the cosmine layer sculptured, but the Glypto-

356 ANNALS NEW YORK ACADEMY OF SCIENCES

pomidz offer a transitional siage leading back to the simple rhombie type,

since in them the scales, although sculptured, are more or less rhomboidal

and overlap but slightly. The Holoptychiidz moreover have excessively

specialized dentrodoni teeth.

The Osteolepide and Rhizodontide are pike-like fishes (Fig. 7), with

elongate bodies and powerful caudal, dorsal and anal fins, which they may

have used both in sudden rushes at their prey and in holding themselves

sill.

Fie. 7.

Restoration oj Megatichithys. Ajier E. D. Weilburn

A progressive osteolepid rhipidistian of the Coal Measures.

Skull-roof: E. cthmoid: Na, nasal; W. nostril; AF. “anterior fronial” [ectethmoid,

septomaxillary]: Fr, frontal; PF, postfrontal: Pa. parietal: S7T.ST, “supratemporals”

[dermosupraoccipital, tabulare]-

Orbital series: AO, “anierior orbital” [lacrymal] ; SO.80, “suborbiials” [jugal]; PO,

postorbiiaL

Temporal series: X. “cheek plate” [anierior portion of true squamosal]; X’, “cheek

plate” [quadratojugal] ; SQ. “squamosal” [supratemporal. pterotic]: P.O. preoperculum

[posterior portion of true squamosal).

Opercular series: Op. operculum: 0, suboperculum.

Gular series: Lj.Lj, lateral jugular [lateral gulars]: the anterolateral gulars may

perhaps give rise to the angulare oi Tetrapoda; J. gular: 4j. “anterior jugular” median

gular.

Gnathal series: Pmz. premaxillary: Ma. maxillary: D. dentary: 1D.1D, infradentaries

[splenial, preangular]; AG, angular [ 7 supraangular].

Shoulder girdle: J.Ci, “iniraclavicular™ [clavicle]; Cl. “clavicle” [cleithrum]; SCL,

“supraciavicle” [supracleithrum].

Fins: B.S. basal scales; RS, ridge scales: FS, fuleral scales; P.Pi 8, propterygial

seales: M.Pi 8, metapterygial scales; AS, anal scales.

Adaptation io predatory habits is indicated by the wide gape and strong sharp teeth,

and by the pikelike body, with powerful median and pelvic fins, which are placed near

the posterior end. The broad. stout pectoral fins would be useful chiefly im suddenly

checking a forward rush and turning the head.

The Holoptychiide on the contrary were relatively deep-bodied, carp-

like forms, with weaker, longer and more flexible fins. The related

Glyptopomidz were more primitive In having rhombic scales and were

pike-like in form, with powerful caudal, dorsals, anals and pelvies.

What were the habits and form of the common ancestors of all these

families? Probably they were comparatively short-bodied forms, with

smal] granular rhombic cosmoid scales, two dorsal fins, a heterocercal

caudal, short lobate pectorals and barely lobate pelvics; the lepidétrichs

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 357

of all fins were short and scale-like; the teeth were conical, but of small

size. In general appearance, except for the shorter body, these primitive

Rhipidistia may have been similar to Osteolepis.

Tarrasius problematicus Traquair (1881, 1890) from the Lower Car-

boniferous of Scotland is usually referred to the Crossopterygii and placed

before the Rhipidistia in a separate order or suborder Haplistia. It is

a very small fish (about 3 inches long) with a continuous dorso-caudo-

anal fin and obtusely lobate pectorals; at least the posterior part of the

body is covered with small granular rhombic scales; the anterior region

of the trunk may have been naked; the notochord is persistent; there are

neural and hemal arches; the median fins are supported by rods, which

are more numerous than the vertebral arches. TYarrasius was provision-

ally referred by Zittel to the Dipnoi, but Traquair (1890, p. 494) said

that “the obtusely lobate character of the pectoral fin seems to point

toward the Crossopterygil.” Goodrich (1909, p. 284) states that the

“dermal bones of the skull and operculum appear on the whole to re-,

semble those of Osteolepids.”

The interest of this fish in the present connection lies in its continuous

median fin. By the upholders of the “‘fin-fold” theory it is assumed that

ancestral fishes once had both the paired and the median fins continuous.

But in the Actinopterygii continuous median fins are invariably a sign

of aberrant specialization; and the frequently copied reconstruction of.a

long-bodied “primitive” fish with a continuous fin-fold bears a suspicious

resemblance to such highly specialized types as Fierasfer, Murena or

Gymnotus. Tarrassius may or may not be a crossopterygian ; but I find

no evidence for believing that its continuous dorsal became subdivided

into the two dorsals of Rhipidistia.

The point is that while paired fins and median fins probably arose in

the same way, they may well have been purely local outgrowths like the

dorsal fin of ostracoderms. It is not necessary to conclude that the two

dorsal fins of Rhipidistia and the single dorsal fin of primitive Actino-

pterygii arose by subdivision or abbreviation of an originally continuous

dorso-anal fin. The Actinopterygii, Rhipidistia and Dipnoi may well

represent parallel offshoots from primitive short-bodied gnathostomes

that had no continuous dorso-anal fin at all, but only low outgrowths of

the skin or of the body-wall, placed at nodal points of mechanical ad-

vantage in securing a “purchase” against the water.

Tarrasius problematicus is also of special interest because, if it is a

primitive crossopterygian, it may also be related to the stem of the

Tetrapoda, retaining perhaps the lobate fins that gave rise to the cheirop-

tergyla, retaining in part a primitive granular scalation, and developing

358 ANNALS NEW YORK ACADEMY OF SCIENCES

a diphycercal tail without powerful rays, such as might be found in an

ancestor of the branchiosaurs.

The endoskeleton of the pectoral limb of the Rhipidistia offers the

only remote approach to the tetrapod type hitherto known among recent

or fossil types. As far back as 1843 James Hall described the pectoral

limb of a large fish from the Catskill formation (Upper Devonian) of

Blossburgh, Pennsylvania, and named it Sauripterus in allusion to the

“sauroid” form of the limb. In 1908 my colleague Dr. Hussakof pub-

lished a small photograph of this specimen in his Catalogue of Types

and Figured Specimens of Fossil Vertebrates in the American Museum

of Natural History, Part I, Fishes (p. 59). As I had long been par-

ticularly interested in the problem of the origin of the Tetrapoda I was

impressed by the almost stegocephalian “look” of the pectoral limb of

Sauripterus, and in February, 1911, I read a paper before the New York

Academy of Sciences entitled “The Limbs of Eryops and the Origin of

Paired Limbs from Fins” (1911) in which I proposed to homologize the

ascending blade of the shoulder-girdle of Sauripterus with the scapu-

locoracoid of Hryops, the single basal element of the fin with the humerus,

the two following elements with the radius and ulna and the remaining

osseous pieces with the carpus and digits. In September, 1912, my

friend Dr. Bertram G. Smith (1912, pp. 540-547) prefaced his excellent

discussion of the phylogeny of the urodeles with a summary of my

Columbia University lectures on the origin of the Amphibia, in which a

Sauripterus-like type of pectoral was assumed as the starting point for

the cheiropterygium; a sketch of the Sauripterus pectoral, by Dr. Hus-

sakof, served to illustrate the subject. In the same year, Professor Patten,

in his book “The Evolution of the Vertebrates and their Kin” (p. 390),

reproduced a photograph of an excellently preserved pectoral limb of

Eusthenopteron (Fig. 8) and said: “Within the pectoral fins, for the

first time in the phylogeny of the vertebrates, appears an axial skeleton

that approaches, in the arrangement of its elements, the characteristic

structure of the appendages of the land vertebrates, 7. e., Husthenopteron.”

In his diagram he homologized the various elements of the fin with the

humerus, radius, ulna, carpus, metacarpus and digits, in much the same

way as I had done in the case of Sauripterus. Early in 1913, Mr. D. M.

S. Watson published in the Anatomischer Anzeiger a note “On the Prim-

itive Tetrapod Limb” in which he also took the Husthenopteron pectoral

as a starting pomt for the cheiropterygium and regarded it as a “reduced

archiptervgium.” In September, 1913, Dr. Broom in his paper “On the

Origin of the Cheiropterygium” gave sketches of the pectoral paddles

of Husthenopteron and Sauripterus and of the shoulder-girdle of the

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 359

latter; he called attention to the resemblances in the teeth between

Sauripterus and Riizodopsis and put forth the hypothesis that the

cheiropterygium had been evolved from the preaxial part of a pre-

Saurvpterus fin. Later in the same year (1913) Dr. Broom published

in the Anatomischer Anzeiger a partial restoration of the shoulder-girdle

aS Se

Fic. 8.—Right pectoral limb of Eusthenopteron foordi. After Patten

A Carboniferous rhizodont rhipidistian.

The broad ascending blade is the cleithrum; the single proximal piece was compared

by Professor Patten with the tetrapod humerus, the next two elements with the radius

and ulna, and the more distal elements with the carpus and digits, respectively.

The single proximal piece (humerus) probably represents the mesopterygium of sharks:

together with the central line of pieces it appears to be homologous with the mesaxial

series in Ceratodus; the divergent rods above and below this axis represent the pre- and

post-axial radials, respectively. The lower main branch is the metapterygium.

and bones of the pectoral paddle of Sawripterus together with a sketch

of the back part of the jaw.

With the view of testing my first idea that the ascending blade of the

Sauripterus shoulder-girdle is the homologue of the scapula of tetrapods

notwithstanding its dermal origin, I have studied the pectoral limb in

various fossil and recent fishes and amphibians. Through the kindness

of Mr. Grimshaw of the Royal Scottish Museum of Edinburgh I was

360 ANNALS NEW YORK ACADEMY OF SCIENCES

permitted to examine the specimens of Megalichthys hibbertu and other

Rhipidistia in that Museum, while in the American Museum I have had

excellent specimens of Osteolepis, Megalichthys nitidus, Sauripterus and

much other recent and fossil material.

In view of the importance of the types specimen of Sauripterus taylors

Hall and of the fact that it has never before been adequately figured, I

have carefully studied this specimen, with the collaboration of my friend

Professor L. A. Adams, whose drawing is here reproduced (Plate I). In

this drawing we have endeavored to show only the structural details that

Vie. 9.—kRight pectoral limb of Sauripterus taylori—Restoration, medial aspect

An Upper Devonian rhizodont rhipidistian, ‘The known parts are shown in Plate I;

the remaining parts are restored from the allied genera J'ristichopterus and Eusthenop-

teron, except the coracoscapula, which is known only from the articular portion and is

conjecturally restored from analogy with Ceratodus.

The mesopterygial series, homologous with the central axis of the “archipterygium”

of dipnoans, Holoptychiide, etc., is represented by the humerus, ulna and distally suc-

ceeding elements; the preaxial radials are barely represented; the postaxial elements

form the major part of the fin skeleton and converge toward the mesopterygial axis.

we have actually seen, without attempting to restore any missing parts.

Areas where the bone has been flaked off, leaving a clear impression in

the matrix, are stippled; cracks and other adventitious features are

omitted. Unfortunately most of the scapulocoracoid, the clavicle (‘“‘in-

fraclavicle”) and the greater part of the fin-rays are missing, and hence

we do not know their precise shape, but have nevertheless attempted a

provisional restoration (Fig. 9) restoring the missing parts chiefly from

the allied genera T'ristichop'erus and Megalichthys. 'The fleshy, scaly

lobe of the fin is indicated by the heavy, curved line. It is not known

whether the ascending bar of the clavicle was present as in Rhizodus or

absent as in Strepsodus (A. 8. Woodward, 1891). We also give for com-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA — 361

parison with Sauripterus a figure of an American Museum specimen re-

ferred to Osteolepis microlepidotus, showing the head, shoulder-girdle

and pectoral fin (Fig. 10).

Further consideration compelled me (1912, p. 220) to give up the

idea that the ascending blade of the shoulder-girdle in Rhipidistia repre-

sents the scapulocoracoid of ‘l'etrapoda, partly because the type of Saurip-

ferus taylori retains a portion of an element that lies between the ascend-

ing blade and the “humerus” or single proximal piece. The rest of this

element probably covered a part of the medial surface of the dermal

ascending blade and may have been present in the counterpart of the

Fic. 10.—Head and pectoral limb of Osteolepis microlepidotus

Crushed specimen in the American Museum (No. 7715). Slightly less than natural

size. The massive cleithrum is partly overlapped by the operculum and suboperculum.

The pectoral fin shows the large scales or plates covering the “humerus,” the elongate

plate covering the postaxial element, or “radius,” the scales on the fleshy part of the

fin and the dermal rays.

type, which was not obtained. In Ceratodus (Fig. 5) a corresponding

piece is apparently homologous with the scapulocoracoid element of tetra-

pods, which is of course a cartilage bone. I therefore return to Gegen-

baur’s view (1895), which is also that of Dr. Broom, that the ascending

blade in Sauripterus represents the cleithrum of primitive Stegocephali,

although in them it is much reduced and the scapulocoracoid correspond-

ingly enlarged. Accordingly the homologies of the elements of the

pectoral limb in Rhipidistia and primitive Tetrapoda (Fig. 11) appear

to be as follows:

RHIPIDISTIA TETRAPODA

Interclavicular corium Interclavicle

Clavicle (“infraclavicle’” ) Clavicle

Supracleithrum (“supraclavicle” ) ?Fused with top of cleithrum (cf.

Sclerocephalus)

Post-temporal Lost

Coracoscapula Coracoscapula

362 ANNALS NEW YORK ACADEMY OF SCIENCES

Flieshy lobe of fin Arm and hand

Mesopterygium (single basal piece) Humerus

Mesopterygial axis Humerus. ulna. ulnare. digit V (7?)

Preaxial parameres (radials) reduced Lost

or absent

Postaxial parameres Radius, carpus, digits I-IV (7?)

Dermal rays Lost

The pectoral limb of Sauripterus differs from the tetrapod type in the

following characters: (1) In Sauripterus, as in other Rhipidistia, the

chief ascending blade of the shoulder-girdle is formed by the cleithrum,

the scapulocoracoid is small and the articular region is protuberant;

whereas even in the most ancient known Tetrapoda the cleithrum is of

relatively small size, the chief ascending blade is formed by the scapulo-

coracoid, and the articular region is sunk below the plane of the sur-

rounding parts. (2) The dermal rays (lepidotrichia) are well developed

(lost in Tetrapoda; cf., modern Dipnoi). (3) The limb is articulated

more on the postero-interior border of the ascending blade than on its

outer face, and is directed backward rather than outward and downward.

(4) The supposed radius, ulna, carpals and digits differ widely in form

from these elements in the Amphibia, so that their supposed homologies

are only recognizable after close study. (5) The successive segments of

the limb apparently were not sharply inclined to each other, as at the

elbow and wrist of Tetrapoda, but were arranged radially more like the

axonosts and baseosts of ordinary fins. (6) While the evidence is not -

positive, the distal rods seem to dichotomize and even the undivided

radials exceed in number the digits of the tetrapod manus. (7) The fin

aS a whole conforms to the imperfect archipterygial type seen in Mega-

lichthys and Eusthenopteron. rather than to the cheiropterygial type of

tetrapods. While all these differences may be simply primitive char-

acters, separating members of two distinct classes, nevertheless they

raise the question whether the tetrapod resemblances in the pectoral fin

of rhizodonts may not be fortuitous, and without phylogenetic signifi-

cance; but in view of the differences in function of the Saurtpterus

paddle and a irue cheiropterygium such underlying similarities as may

exist can hardly be attributed to convergence, and when taken in con-

nection with the resemblances in the skull above noted (pp. 332-337)

they gain in importance.

Dr. Broom’s view that the tetrapod cheiropterygium developed only

from the anteroventral border of a pre-Sauripterus stage appears to me

to lack adequate evidence. Nor do I consider the presence of dermal

rays as unfavorable to the development of a cheiropterygium. The re-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 363

Fie. 11.—Comparison of the paired limbs of Paleozoic rhizodonts and tetrapods

Left pectoral limbs of (A) Sawripteruws and (B) Eryops. Medial or palmar surface.

Left pelvic limbs of (C) Husthenopteron, after Goodrich, and (D) ZHosauravus, after

- Williston. Medial surface.

The elements that converge toward the mesopterygial axis (ulna, fibula) are hatched.

364 ANNALS NEW YORK ACADEMY OF SCIENCES

duction of the dermal rays has frequently occurred in the Actinopterygii

(e. g., Murena, Lepidosiren) without impairing the locomotive power of

the fish. When the pectoral fin was turned downward and forward the

medial surface of the spreading lobate portion of the fin would seem to

be fitted for the digital emargination of the borders and the modification

of the radials to form the digits. Dr. H. H. Wilder (1909, p. 235) has

also called attention to the fact that in the manus of Necturus the

muscles for spreading and closing the digits are so highly developed as

to suggest derivation from a primitive appendage which had the power

of widely spreading and closing, as in the fins of fish. In Sauripterus

(Fig. 10) and Husthenopteron (Fig. 8) most of the radials converge

toward the mesopterygial axis, which includes the supposed homologues

of the humerus, ulnar, ulnare and digit V (?), and the same is true of

the digits of the manus of Carboniferous Amphibia (Fig. 11) and Rep-

tilia and of recent urodeles; in both classes the post-axial paramere, or

radius, is sharply separated from the remainder of the cheiropterygium.

I find that both Emery (1897, p. 208) and Jaekel (1909) have also

noted this biramous character of the tetrapod cheiropterygium.

The muscles of the pectoral limbs of Sauripterus were probably sep-

arable into a deep proximal mass covering the scapulocoracoid and the

humerus and a deep distal mass running from the humerus to the distal

radials. Surface muscles perhaps extended from the radials back to the

scapulocoracoid region. The joint corresponding to the elbow joint is

clearly present in these rhizodonts and the “‘radius” and “ulna” are seen

to be a part of the outer or distal segment of the limb.

The pelvic limb of Husthenopteron (figured by Goodrich, 1909, p. 275),

which is the best known one of the rhizodonts, has likewise a certain re-

semblance to the tetrapod type, as noted by Jaekel (1909), in so far as

it possesses a single basal piece analogous to the femur and two main

radials analogous to the tibia and fibula. The pelvic limb, lke the

pectoral, appears to represent an imperfectly attained mesorhachic type

in which the preaxial elements have become more or less produced later-

ally and to some extent regrouped. The “femur” has two broad distal

facets for the supposed fibula and tibia. The mesopterygial element,

which may be called the “fibula,” is a wide element bearing at its distal

end two facets for the proximal “tarsals” which appear to be the fibulare

and intermedium, or tibiale. The distal end of the supposed tibia, or

preaxial paramere, lies beside the supposed intermedium; all the tarsals

slant toward the fibular or mesopterygial axis. The joints corresponding

to the knee and ankle joints are well defined (Fig. 11). All this con-

stitutes a distinct resemblance to the pelvic limbs of Carboniferous Tetra-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 365

poda (cf. Jaekel, 1909, Figs. 2-18), the great difference being that in the

tetrapod the tarsals have become relatively shortened and wider while the

_ metatarsals and digits, the tibia and femur have lengthened.

The stout pelvic bones appear to represent the ischio-pubis. No

known representative of the ilium is present in the rhizodonts; but even

in such relatively advanced tetrapods as Hryops, although the ilium is

very large, it has not yet gained a secure contact with the backbone, and

we must either suppose that the dorsal growth of the pelvic cartilage

followed very rapidly upon the change in function from fins into support-

ing limbs; or possibly that the anterior process of the “ischio-pubis” of

Eusthenopteron became rotated upward and gave rise to the ilium, while

the posterior expanded portion broadened out into the true ischiopubic

mass.

STEGOCEPHALI

Between the oldest known Amphibia of the Coal Measures and all fishes,

there remain profound structural differences which are as yet unbridged

by paleontological discovery. Even the branchiosaurs and their still

more degenerate modern successors the urodeles have the dermal shoul-

der-girdle reduced and the epiphyses of their limb-bones cartilaginous,

which may indicate that instead of being primarily aquatic animals, true

links between fishes and terrestrial quadrupeds, they are secondarily

aquatic (see also B. G. Smith, 1912, pp. 547-551). And yet in the larval

state the branchiosaurs undoubtedly retained true piscine branchial

arches, while the branchial skeleton of the Permocarboniferous “urodele”

Lysorophus, as figured by Williston (1908), can leave no doubt of an

ultimate piscine origin.

So wide are the differences between the various groups of Paleozoic

Amphibia that one is led at first to inquire whether they may not have

come off from different, but allied, groups of fishes, so that for example

the Steresopondyli and the Temnospondyli might be related to the Osteo-

lepide and Rhizodontide, while the Branchiosauria, Microsauria, Aisto-

poda and Urodela might conceivably run back to some form like Tarras-

sius (see pp. 357-358 above). Yet in spite of these wide differences

among the Paleozoic Amphibia they all agree in having one fundamen-

tally identical skull pattern and cheiropterygial limbs, so that present

evidence suggests that the transformation of fishes into amphibians oc-

curred but once.

This transformation involved first of all the abandonment of the tail

as the principal propeller, the loss of its-dermal rays, the abortion of the

hypural bones and the assumption of the gephyrocercal form. Such a

366 ANNALS NEW YORK ACADEMY OF SCIENCES

transformation has occurred many times in the Actinopterygii and also

in the Dipnoi. If the pro-Tetrapoda were Rhipidistia the transformation

also involved the loss of the two dorsal and anal fins and of their ex-

panded basal supports. The notochord in these pro-Tetrapoda must have

been wholly persistent, for in the earliest known tetrapods the vertebrz

are not only of widely varied type, but by no means complete. The pro-

Tetrapoda undoubtedly had both the pectoral and pelvic fins strongly

developed and of lobate or fleshy-based type. When they first emerged

from the water, either in pursuit of littoral prey or during times of

drought, they may have used the medial or palmar surface of the paddles

for traction and propulsion, at the same time wriggling on their bellies.

Some of those with small paddles soon adopted a snake-like habitus and

lost the hmbs (Aistopoda, Apoda). In those that developed the limbs

(Fig. 14) the scapulocoracoid increased rapidly im size and became ossi-

fied, the cleithrum was reduced, the posttemporal and supracleithrum

disappeared and the only connection between skull and shoulder-girdle

was furnished by the “trapezius” muscle, which in modern urodeles ex-

tends from the “suprascapula” (? cleithrum) to the occiput (Fig. 14, F).

Thus the shoulder-girdle acquired mobility, while, with the growth of the

scapulocoracoid, its muscles acquired a larger base, and extended around

from the medial to the external side. The extension of the coracoscapular

muscles, ventrally, dorsally and externally, together with the differentia-

tion of the pectoral muscles greatly increased the strength of the forearm

and crowded the cleithrum to the front edge of the scapula. The differ-

entiation of the pectoral muscles conditionea the formation of the rhom-

boid interclavicle which partly overlies them. F

More difficult to comprehend are the muscular readjustments which

must have ensued when the elbow and knee bends were being established,

and the shifting of the articular surfaces of the humerus radius and ulna.

With regard to the muscles of the cheiropterygium itself, Professor

H. H. Wilder (1909, pp. 230-231, 235) im the course of an illuminating

discussion of the musculature of Nectwrus writes as follows:

“The muscles of the distal portion of the vertebrate chiropterygium, that is,

from elbow or knee on, aside from the modifications imposed upon them by

the varying shapes of the limbs themselves, and the great difference in their

use, are, in their essential features, quite similar in all living forms, and in

their differences show the modifications of a primary type due to environment

rather than the suggestions of an historic development of that type. The

study is, therefore, one mainly of the adaptations of a given set of elements,

rather than a phylogenetic history, which latter, as is the case also with the

bones of the same region, must be sought in the gap separating fin and hand,

that is, in the phylogenetic stages represented by lost forms of ganoids, stego-

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 367

cephali, and their allies. The salamander Necturus, probably the nearest ap-

proach to this series represented by living fauna, offers in its distal muscles

some few suggestions of an earlier phylogenetic stage, and is thus of funda-

mental importance in the present inquiry. The well-nigh complete correspond-

ence in the fore and hind limb as regards not only bones and muscles, but

other parts as well, has been commented on above and offers strong support

for the doctrine of serial homology, to be considered later. There are, also, as

is the case with higher forms, some traces of a correspondence between the

dorsal and ventral surfaces of a single paw, giving a suggestion of the deriva-

tion of the chiridial musculature from a fin-like precursor in which the jointed

rays (digits) were supplied by similar muscular elements applied both dor-

sally and ventrally, as in present-day fishes. The following description is that

of the anterior limb, but with the substitution of the terms tibia and fibula

for radius and ulna, tarsus for carpus, and so on, it will be found almost

equally applicable to the posterior one. In a few cases a muscle which is well

developed in the anterior limb is small or wanting in the posterior, and thus

the former is a little more typical.’

“Reviewing the conditions in this, probably the most primitive chiroptery-

gium now left to us, several interesting points become manifest. The digits

are moved in two ways, either flexed and extended or moved sideways, but

while the system which provides for this latter form of motion is extremely

well perfected, that for flexion and extension is not. For abduction and ad-

duction there are typically five separate muscles for each digit, that is, two

ventral, two dorsal and one intermetacarpal, while for flexion and extension,

aside from the system supplied by an aponeurosis, and evidently a newly in-

troduced feature, there are but three. This extreme perfection of the sideways

movement of the digits in the most primitive chiridium known, together with

the weak and makeshift arrangements for bending and straightening the digits,

strongly suggest the derivation of the chiridial type from one in which the

digits (fin-rays?) required to be constantly opened and shut by lateral move-

ments, precisely as in the case of the fins of most fishes.

“During later phylogenetic history there is an evident tendency to increase

the efficiency of the flexor-extensor system and diminish that of the abductors

and adductors, except in the case of the two digits that form the ends of the

series (I and V), and the most of these changes have already occurred among

the higher urodeles.”’ .

As the skeletal remains of the limbs of Carboniferous Tetrapoda retain

but little that is clearly suggestive of derivation from the paired appen-

dages of fish, so too the footprints of these animals indicate that the

8 “Tn one point the free limb of Nectwrus diverges from what is generally believed to

be the typical chiropterygium, and that is, it possesses but four digits in each extremity

instead of the canonical five which is usually considered primitive. Since the nearest

ally of this species, the cave form, Proteus, exhibits a still greater reduction of digits

(anterior, 3; posterior, 2), it has been presumed that this is in both cases a secondary

reduction. Certain facts, however, lead one to think that the first land vertebrates pos-

sessed a smaller number of digits than five, and if this be so, the condition in these two

salamanders is primitive. and not a secondary reduction. According to the reduction

theory digit I is assumed to be the one lost, and in accordance with this the four digits

present are designated here. both in text and illustrations, as II-V.” (Wilder, p. 251).

368 ANNALS NEW YORK ACADEMY OF SCIENCES

fleshy portions of the limbs were perhaps no more fish-like than in the

modern Necturus.

These Carboniferous footprints vary widely in form, as do also the

known skeletons of Carboniferous Tetrapoda. The number of toes in

both manus and pes varies from three to five (G. F. Matthew, 1903,

1904), but in perhaps the majority of cases the manus has four digits

and the pes five. Abel (1912, p. 68) indeed concludes that no stego-

cephahan had a five-fingered hand; but in a well known specimen of

Eryops megacephalus (Amer. Mus., No. 4186) which was described by

Cope (1880) digits I, III, IV, V are present in the fossil and the missing

digit II is represented by a wide facet on carpal 2.

One of the oldest known footprints is a single impression from the

Mauch Chunk Shale (Lower Carboniferous) of Pennsylvania to which

the name Thinopus antiquus was given by Marsh (1896). ‘The imprint

consists of two stout, jointed toe-marks, which are nearly parallel to each

other, but separated by a considerable interval extending back to the mid-

dle of the palm; from one of the toes a smaller offshoot near the tip indi-

cates a small lateral toe. It may be a mere accidental resemblance that

the pes of the modern Proteus is likewise bilobate, even in early stages

of development (Wiedersheim, 1892, p. 199). A footprint named Asperi-

pes avipes from the Carboniferous of Eastern Canada (G. F. Matthew.

1904, Pl. II, fig. 2a, 2b) represents a three-toed manus that may have

been somewhat similar to that of Thinopus antiquus; but the pes of

Asperipes has five digits.

Thus it is an open question whether the three- and four-toed feet of

Carboniferous Tetrapoda represent reduction stages from the typical

pendactyl cheiropterygium, or whether there has been an increase in the

number of the digits from three to five. Favoring the former supposition

is the following evidence: In modern Salamanders the number of digits

is four in the manus and five to four in the pes; in the pes the process

appears to have been reduction from five to four rather than the reverse:

for, (a) in the development of the pes of the four-toed Salamandrella

kayslerlingii there appears a vestigial fifth tarsal which later unites with

tarsal 4 (Schmalhausen, 1910, figs. 6, 7), and (b) in the degenerate

Proteus the number of digits is reduced to three in the manus and two

in the pes. .

In many of the Carboniferous footprints the fifth or outer digit of both

manus and pes is sharply divergent, the fourth is the longest and the ends

of digits IV, III, II, I are turned inward, while the foot as a whole points

forward; all these characters suggest limb-structure fundamentally simi-

lar to that of Eryops or indeed of modern urodeles, especially the sala-

manders.

~~.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 369

An important fact in connection with the origin of the tetrapod limbs

is that in all known Carboniferous Amphibia and Reptilia as well as in

recent urodeles all the digits, together with their carpals or tarsals, con-

verge toward the ulna or fibula respectively, and that the whole carpus

and tarsus are obliquely placed with reference to the ulna and fibula.

This arrangement appears to be foreshadowed in the rhizodonts (Fig.

11) ; especially if we assume that only those radials that converge toward

the mesopterygium were preserved, and that the postaxial radials gave

rise to the vestigial prehallux and prepollex which are so widely distrib-

uted in the Tetrapoda (cf. Emery, 1897).

Although an ilium and a sacral attachment to the backbone are not

known in fish, the readiness with which the backbone forms an attach-

ment with the bases of the fins in different groups of fishes, either through

the hypural bones, or, in the Rhipidistia, with the expanded bases of the

dorsal and anal fins, indicates that with the enlargement of the hind

hmbs and pelvis the development of paired dorsal apophyses or ilia and

their subsequent attachment to the sacral ribs would soon follow. Indeed

the sacral ribs of Hryops and even of modern urodeles furnish an exam-

ple of the imperfect connection between the pelvis and the backbone that

existed in early tetrapods.

“PROCORACOID AND CORACOID” OR “CORACOID AND METACORACOID” ?

Notwithstanding all that has been written since Howes’s paper (1887)

in the effort to homologize the coracoidal elements throughout the Tetra-

poda, investigators are still divided as to the application of these terms

and as to the implied homologies. Professor Williston, in various publi-

cations, inclines to the view that the true mammalian coracoid is repre-

sented in the primitive reptiles by the anterior one of the two coracoidal

elements ; he accordingly calls it “coracoid,”’ while the posterior element

he calls “metacoracoid” ; other investigators, however (including Huene,

Broom and Watson), continue to use the terms “procoracoid”’ and “cora-

coid.”

With the hope of coming to some decision in the matter I have com-

pared the shoulder-girdle of representative Tetrapoda of all classes and

Professor L. A. Adams has prepared the series of figures (Figs. 12-15)

here published. It seems to us that Broom’s observations (1899) on the

development of the shoulder-girdle of marsupials offer decisive evidence

that the true coracoid is the main element that extends down to the

sternum in embryonic marsupials (Fig. 12, A, B), but we regard the

mesenchymatous anterior element called by Broom “procoracoid” as more

probably homologous with the epicoracoid of the monotremes and lizards

TO ANNALS NEW YORK ACADEMY OF SCIENCES

Fic. 12.—Homology of the coracoid in primitive mammals

A. Left shoulder-girdle (outer view) of an 8.5 mm. embryo of a diprotodont marsupial,

Trichosurus vulpecula. After Broom. In this stage the true coracoid (cf. figs. B, C)

is connected below with the sternum, as in adult monotremes. The epicoracoid (“‘pro-

coracoid’” of Broom) is mesenchymatous. It does not share in the glenoid and is there-

fore unlike the reptilian “‘procoracoid’ (—coracoid), but does resemble the membranous

epicoracoid.

B. Left shoulder-girdle (ventro-lateral view) of a 14.8 mm. Trichosurus vulpecula

embryo. After Broom. The coracoid at this stage retains its connection with the

sternum, which connection is lost in the adult: the epicoracoid has disappeared.

C. Right shoulder-girdle and proximal part of humerus (inner view) of adult diproto-

dont marsupial, Phascolarctus cinereus. The single coracoid shares in the glenoid and

points downward and inward toward the sternum.

D. Left scapulocoracoid (vyentro-medial view) of Dasypus sp., showing the metacora-

coid (m.co) as a separate element.

E. Lower part of left scapulo-coracoid of Megatherium (inner view), after Weber,

showing suture between coracoid and metacoracoid.

F. Left scapulo-coracoid of Lepus (outer and distal views), after Howes, showing

coracoid and metacoracoid both sharing in the glenoid articular surface.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 371

(see below), and we do not favor Broom’s view that the posterior, glenoid-

sharing element (Fig. 12, D, E, F) in rodents, edentates and many other

Fig. 13.—Homology of the coracoids in Monotremes

A, Shoulder-girdle of Echidna aculeata; B, Ornithorhynchus anatinus 3 left side, outer

view; C,D, the same, ventral view.

The epicoracoid (€p.co) agrees with that of lizards and Sphenodon in all its relations.

The broad coracoid may either represent both the primitive reptilian eiements (coracoid

and metacoracoid) or the metacoracoid may have disappeared.

placental mammals is a neomorph, im the nature of a glenoid epiphysis.

This so-called epiphysis agrees with the “metacoracoid” (Williston) of

Permian reptiles (Fig. 14, E, F) in its relations with the true coracoid

372 ANNALS NEW YORK ACADEMY OF SCIENCES

mop

Fic. 14.—Homologies of the coracoids, etc., in reptiles

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 373

A, Sphenodon, right shoulder-girdle, outer view; B, ditto, ventral view. The coracoid

is single. The metacoracoid may have disappeared. The epicoracoid (ep.co.) as in the

lizard (C,D) is bounded by the scapula, coracoid, interclavicle and clavicle. Beneath the

T-shaped interclavicle is the intercoracoid, or anterior sternal region.

C, Varanus, left shoulder-girdle, outer view; D, ditto, ventral view.

In Sphenodon and Varanus the so-called “‘suprascapula”’ (? cl) may possibly represent

the cleithrum of primitive Tetrapoda (cf. Fig. 14 D). The ‘“‘suprascapula,” like the

cleithrum, is connected below with the clavicle; to it is attached the trapezius muscle.

The coracoid is single; anteriorly it is fenestrated for the pectoral muscles.

BH, Shoulder-girdle of a Cotylosaur, Labidosaurus. Ventral view, after Williston. The

coracoid and metacoid are distinct; the cleithrum is absent.

F, Shoulder-girdle of a Permocarboniferous reptile, Hdaphosaurus novomezicanus,

after Williston and Case. Right side. The expanded coracoid region, short scapula and

large clavicle are perhaps inherited from primitive aoe wars The metacoracoid is

present.

and with the glenoid; its wide distribution as a vestigial or reduced

ossific center in various orders of mammals suggests that it is an ancient

reptilian inheritance.

In reference to the shoulder-girdle of monotremes we would apply the

name coracoid to the posterior element, which forms part of the glenoid

(Fig. 13), since this part closely resembles the true coracoid of embryo

marsupials. The anterior element (Fig. 13, ep. co.) may correspond

with the epicoracoid (Fig. 14) of lizards and Sphenodon, as suggested

by Wiedersheim (1909, p. 190), which never participates in the glenoid

and always lies beneath the interclavicle and clavicle. Thus the true

coracoid of monotremes may either have resulted from the loss of the

suture between the coracoid and metacoracoid or the metacoracoid may

have disappeared, while the epicoracoid has been developed, perhaps in

adaptation to fossorial habits.

In the lizards, Sphenodon and other reptiles with a “single” coracoid

(Fig. 14), this singleness may also have resulted either from the loss of

the suture separating the coracoid from the metacoracoid, or from the

disappearance of their coracoid. Their epicoracoid is also well developed

and ossified.

In the lower Tetrapoda (Fig. 15), including various Permian orders,

the coracoid and metacoracoid are often divided by suture, but the epi-

coracoid is not ossified.

In Ceratodus (Fig. 5), the sturgeons (Fig. 4) and the sharks, which

have the best developed and largest coracoscapula cartilages among recent

fishes, there is no sutural separation of parts and it seems likely that the

segregation of ossific centres corresponding to the coracoid and metacora-

coid is a later advance, perhaps correlated with the great expansion of the

coracoscapular mass and the higher differentiation of the limb muscles

in tetrapods.

re ANNALS NEW YORK ACADEMY OF SCIENCES

Fic. 15—Homologies of the coraecoids, etcetera. in Amphibia

A. Shoulder-girdle of a Permocarboniferous temnospondsl. Eryops megacephalus,

outer view of leit side; irom specimens in the American Museum: B, the same, ventral

view: C. the same. ventro-<xtermal view. (No suture between the coracoid and meia-

coraceid was found)

All the elements of the primitive tetrapod shoulder-girdle are present The median

ventral space between the coracoids was probably fjlled by the intercoracoid mesen- ’

chymatous tissue.

Dermal elements: ic. interclavicle : cr. davicle: ci. cleithrum.

Cartilage bones: ¢6. coracoid: m.co, metacoracoid : sc. scapula: gi, glenoid.

GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 375

The expanded upper portion of the cleithrum may have served for the attachment of

the trapezius muscle (cf. Fig. 14, F).

D, Shoulder-girdle of a Permocarboniferous temnospondyl Cacops aspidephorus. After

Williston.

The cleithrum extends over the top of the scapula after the fashion of a suprascapula

(cf. Fig. 14, F). The lower end of the cleithrum may represent the mammalian acro-

mion (Gaudry, Broom).

E, Shoulder-girdle (ventral view) of a modern urodele, Salamandra maculosa, after

W. K. Parker. Nearly adult, enlarged 4/1. The greatly enlarged intercoracoid car-

tilages, which overlap in the mid-line, furnish a broad base for the pectoral muscles.

The anterior extension occupies the position of the epicoracoid. The inierclavicle and

clavicle are lost. st, sternum.

F, Shoulder-girdle (left side, outer view) of Salamandra maculosa, after W. K. Parker.

Nearly adult, enlarged 5/1.

The coracoid, metacoid and scapula are still separated by sutures. The dermal ele-

ments are lost, except possibly the cleithrum (? cl), which may be represented by the

so-called suprascapula; this gives attachment to the trapezius muscle.

G, Shoulder-girdle of a Permocarboniferous branchiosaur, Brachiosaurus salaman-

droides. After Fritsch. Viewed from above. x6/1. In the branchiosaurs, which are

in many characters structurally ancestral to such modern urodeles as Cryptobranchus,

the intercoracoids are much expanded, the scapula has a short truncate blade and the

dermal elements (clavicle, interclavicle, cleithrum) are slender.

H, Scapulocoracoid of a Permocarboniferous reptile, Dimetrodon. After Williston.

The expansion of the coracoid and the small size of the metacoracoid are reminiscent

of the temnospondylous Amphibia. This scapulocoracoid is fundamentally similar to

that of the Therocephalia and doubtless represents a pre-mammalian stage. acr, acromial

process.

In short we conclude: (1) that in primitive reptiles the homologue of

the true mammalian coracoid is that anterior element which: (a) lies

below the acromial border of the scapula, (b) above the clavicle, (c)

enters the glenoid articulation posteriorly and (d) extends ventrally

toward the sternum; (2) that the posterior element or metacoracoid in

many mammals persists in a reduced condition, arising from a separate

center; (3) that in monotremes, lizards, Sphenodon and certain other

reptiles the suture between the coracoid and metacoracoid may have

disappeared or else the metacoracoid has disappeared (Williston) ; (4)

that the epicoracoid is a thin membranous element lying between the

coracoid and the interclavicle in lizards, Sphenodon and the monotremes,

which was probably present also in many of the Permian Tetrapoda.

CONCLUSION

Although the foregoing discussion of the derivation of cheiropterygial

limbs from piscine appendages is regrettably incomplete, it 1s at least

based so far as possible upon the combined data of comparative anatomy

and paleontology, and the evidence here summarized is, I hope, in line

with the historical trend of development in these fields.

From the viewpoint which has been developed in the preceding pages,

the origin of the Tetrapoda is regarded as a remote consequence of the

376 ANNALS NEW YORK ACADEMY OF SCIENCES

transformation of minute, acraniate, radially constructed ccelenterates,

moving by the lashing of cilia, and ingésting microscopic food particles,

into pre-gnathostome animals with contractile myocelomic pouches,

which, becoming muscular, gave rise to the primary locomotive organs,

or myomeres, as well as to the ccelom.

Bilaterality ensuing, and pharangeal diverticula appearing, our ani-

mals entered the path of predatory adaptation; this led to the acquire-

ment of accessory locomotive structures, including the local outgrowths

called fins, and to the formation of a synthetic or gnathostome type of

head, fitted for seizing and ingesting living prey. .

The acquisition of a many-layered skin, proceeding with the further

differentiation of the myoccelomic pouches, and of the derived mesen-

chyme, was followed by the development of an exoskeleton, and later of

an endoskeleton, the mesenchyme carrying the bone cells either to the

skin or to the mesenchymal connective tissue sheaths between the myo-

meres and around the dorsal axis.

Endoskeletal supports thus evolving the Osteichthyes appeared.

Of these the Dipnoi acquired elaborate tritoral teeth and other special-

ized conditions and thus removed themselves from the main ascending

line, while in the other direction the Actinopterygii carried to a high

perfection the scale-like dermal rays and other purely aquatic adaptations.

The Crossopterygii partly paralleled the Actinopterygii in the evolu-

tion of dermal rays but were distinguished by the outgrowth of fleshy

lobate, fan-like paired fins, the prerequisites for the acquirement of hands

and feet. The double-breathing pro-Tetrapoda reversed the direction of

their evolution, sacrificed dermal rays, hypural bones, caudal and dorsal

fins, scales, operculars, gulars and the elements connecting the shoulder-

girdle with the skull, largely reduced the cleithrum and emerged from

the water by virtue of the tractive and propulsive power of their stout

paired fins.

As thus conceived the rise of the vertebrates and the origin of the

Tetrapoda constitute a history of successive improvements in the loco-

motive apparatus.

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GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 379

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380 ANNALS NEW YORK ACADEMY OF SCIENCES

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Sitzungs. Ber. Gesellsch. Naturforschender Freunde, Nr. 10, Jahrg. 1909,

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as a Memorial of the late John Samuel Budgett, pp. 195-284, pll. xiii-xy.

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Vol. XXX, pp. 167-179. 1881.

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Rocks of Eskdale. Ann. Mag. Nat. Hist., Vol. VI, p. 494. 1890. [Tarrasius

problematicus. |

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382 ANNALS NEW YORK ACADEMY OF SCIENCES

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Trans. Roy. Soc. Edinb., Vol. XLI, Pt. ii, pp. 469-475. 1905.

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pp. 1-28. 1909.

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: Micropholis Stowi, Huxiey,a Temnospondylous Amphibian from South

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: Batrachiderpeton lineatum Hancock & Athey. a Coal Measure Stego-

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: On the Primitive Tetrapod Limb. Anat. Anz., Vol. XLIV, pp. 24-27.

1913.

: Notes on Varanosaurus acutirostris, Broili. Ann. Mag. Nat. Hist.,

Ser. 8, Vol. XIII, pp. 297-310. 1914. [Homologies of the upper and lower

temporal elements, pp. 309-310.]

: Pleurosaurus and the Homologies of the Bones of the Temporal Re-

gion of the Lizard’s Skull. Ann. Mag. Nat. Hist., Ser. 8, Vol. XIV, pp. 84-

95. 1914.

WELLBURN, E. D.: On the Genus Megalichthys, Agassiz: Its History, System-

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Vol. XIV, Pt. I, pp. 52-71. 1900.

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Canada. Trans. Roy. Soc. Canada, Vol. IV, See. IV, Pi. Dl, peewee

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beriicksichtigung des Schulter- und Beckengiirtels bei Fischen, Amphibieu

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: Vergleichende Anatomie der Wirbeltiere, Tth Ed. Jena, 1909.

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1908.

: New or Little Known Permian Vertebrates—Pariotichus. Biol. Bull,

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: New or Little Known Permian Vertebrates—Trematops, new Genus.

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: Cacops, Desmospondylus; New Genera of Permian Vertebrates. Bull.

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i-iii. 1910.

: New Permian Reptiles: Rhachitomous Vertebrex. Jour. of Geol., Vol.

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- American Permian Vertebrates. Uniy. Chicago. 1911.

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GREGORY, PRESENT STATUS OF ORIGIN OF TETRAPODA 383

WILLISTON, S. W.: Restoration of Seymouria baylorensis Broili, an American

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Woopwarp, A. 8.: Catalogue of the Fossil Fishes in the British Museum (Nat-

ural History), Part II. 1891. [Rhizodontidz.]

: Outlines of Vertebrate Palzxontology. 1898. [Comparison of skull-top

of Rhizodopsis and Stegocephalian, pp. 124, 125.]

: The Study of Fossil Fishes. Proc. Geologists’ Assoc., Vol. XIX, pp.

266-282. 1906.

Zwick, W.: Beitrige zur Kenntnis des Baues und der Entwicklung der Am-

phibien Gliedmassen, besonders von Carpus und Tarsus. Zeitschr. f. Wiss.

Zool., 63ter Bd., ss. 62-114, Taf. iv, v. Leipzig, 1898.

RIGHT PECTORAL GIRDLE AND LIMB OF Sauripterus taylori HALL

Type, Amer. Mus. No. 3341. Internal, or medial, view. Two-thirds natural

size. .

el, cleithrum; swu.cl, ? supracleithrum; co.sc, remnant of coracoscapula, ad-

hering to cleithrum and possibly represented also by the small patches p, p’

(Broom) ; ”, notch, possibly for ascending blade of clavicle; S, enlarged scales

on metapterygial border ; 8’, scales on lobate portion of fin; H, single proximal

basal piece, or humerus; U, mesopterygial axonost, or “ulna”; R, metaptery-

gial axonost, or “radius”; r, radials; dr, dermal rays.

a gortodety Mensa ost zarie

gakene hegislns .2 ; a ae ee

VoLuUMn XXVI, PLATE IV

ANNALS N. Y. ACAD

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PRESS OF JUDD & DETWBRILER, INC., WASHINGTON, D.C.

ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXVI, pp. 385-394, pl. V

Editor, E>MuND OTIs Hovey

BIOCHEMICAL STUDIES OF SELENIUM

Victor EK. LEVINE

NEW YORK

PUBLISHED BY THE ACADEMY

18 OcroBEr, 1915 a

THE NEW YORK ACADEMY OF SCIENCES

(LycEumM oF NaTURAL History, 1817-1876)

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[ANNALS N. Y. AcaD. Scr., Vol. XXVI, pp. 385-394, Pl. V. 18 October, 1915]

BIOCHEMICAL STUDIES OF SELENIUM? |

By Victor E.. LEVINE |

(Presented in abstract before the Academy, 18 May, 1914)

CONTENTS

Page

ISMVOONCEION.....:...03 0: eee eae aT et a Side a. abel oe esa v/a, a baa Ole SRM OO aR 385

Sodium selenite as a reagent for reducing substances................000. 386

Reduction of selenium compounds in the living organism................ 388

ae PE HITIS ANICE Se oe. sce cree eee wieco Sle ios a vi wise Rae ae Wsipiae Grerelats oa OS ore 388

(| BS Fra WS AE OU ick Cop A ae 388

OT EDU er aes) Oe fac BLSRTEINS 2 NE 388

INRA MEETS UNOS 50 soi eter Maen gees Wo keh Leia Ai dg. boa w.6 Wi ou OS Rig ute b SEND 389

Selenium compounds injected into the animal system............... 389

TIES PE DPSS Ra ae at oe Se er 2 ea 390

Effect of selenium compounds upon catalase and other enzymes.......... 391

MMM AGE Pee os See sos EWI ERIS a a here lie hl Saat dls Sen ee eee Se 391

CELLU AIST SETS LSS Ce Ie es a een oe ee oe a Pee 392

RS ae oa a ec iar eia AUS Ria, wk Ora S WM ee eR aw We Subang’ 392

2 DEPOSED. 2 2 cy EERSTE Be ean eas 392

aR Ps eRe EIA Nava ph Ske od ale AU's thee eid oe BS wets Ow > 392

Weil PU EAE SV LEUSE 06 pa a ee en 393

uP LL TG SU EC Ea en 393

ee eePeere OF TSClENIUIN COMPOUIGS . 6.0. ec eke cee te eee eee e ec ewen 393

INTRODUCTION

There has been of late an ever increasing interest in the quest for

information concerning the biologic action of the rare elements. Within

the last few years attempts have been made to introduce radium, thorium,

cerium, vanadium, tellurium and others into therapeutics; and it is but

very recently that palladium has been suggested as a cure for obesity and

selenium as a valuable agent in cancer therapy. Furthermore, selenium

has become of increasing interest since it has been found (to the extent

of 0.2 mg. per liter) in the mineral waters of La Roche-Posay, and since

some of its compounds can serve as excellent reagents for the detection

of certain alkaloids and also for the detection of micro-organisms.

The experiments undertaken extend the work, in this laboratory, which

was done on selenium by Woodruff and Gies more than a decade ago.

1 Manuscript received by the Editor 27 March, 1915.

(385)

886 ANNALS NEW YORK ACADEMY OF SCIENCES

The experiments relate to the reduction of selenium compounds hy chem-

ical substances of biologic significance, by micro-organisms, by plant sub-

stances and by animal tissues. ‘Toxicological and pharmacological effects

were also studied, as well as the effect on germination and growth of

plants, the effect on enzyme activity and the effect on the precipitation

of proteins. The compounds employed were selenium dioxid (selenious

acid), sodium hydrogen selenite, normal sodium selenite, selenic acid,

sodium selenate, potassium selenocyanate. For some of the compounds

thanks are due to Professor Victor Lehner, of the University of Wisconsin.

Sopium SELENITE AS A REAGENT FOR REDUCING SUBSTANCES

Sodium selenite in alkaline solution can be used as an indicator for

reducing substances, especially for the carbohydrates containing a free

carbonyl. group. ‘The following do not reduce sodium selenite (alkaline

to litmus) : acetone, formaldehyde, trioxymethylene, acetaldehyde, furol,

benzaldehyde, cinnamic aldehyde, salicyl aldehyde, piperonal, methyl al-

cohol, ethyl alcohol, glycerol, erythrol, mannite, inosite, phenol, the

eresols, thymol, a-naphthol, acetic acid, butyric acid, B-oxybutyric acid,

palmitic acid, stearic acid, trichloracetic acid, oxalic acid, tartaric acid,

citric acid, oleic acid, malic acid, cinnamic acid, hippuric acid, glvcocol,

alanin, guanidin carbonate, leucin, urea, thio-urea, ammonium sulfo-

cyanid, caffein, theobromin, uric acid, sodium urate, creatinin, lecithin,

cholesterol, palmitin, stearin, olein, blood albumen, blood fibrin, edestin,

egg albumen, gelatin, peptone, proteoses, ovalbumin, collagen, osseo-

mucoid, elastin, saccharin, antipyrin, anthraquinone, sucrose, raffinose,

cellulose, starch, dextrin, glycogen, inulin, esculin, amygdalin, and the

following gums: arabic, tragacanth, guaiac, rosin, benzoin, kino, aloes,

asafoetida, myrrh, gambir. Alcoholic solutions of benzoin, kino or aloes

give a red-brown to cherry-red solution without the addition of sodium

selenite. The following reduce sodium selenite: amidol, arabinose, rham-

nose, xylose, dextrose, galactose, levulose, maltose, lactose, hydroquinone,

phloroglucin, pyrogallol, hydroxylamin hydrochlorid, phenylhydrazin hy-

drochlorid, benzidin hydrochlorid, hydrazin hydrate; arsenious, hydro-

bromic, hydriodic, phosphorous, hypophosphorous and sulfurous acids;

ferrous sulfate, stannous chlorid, zine and hydrochloric acid, hydrogen

sulfid, acetylene, formic acid, gallic acid, lactic acid, tannic acid.

Acetone, acetaldehyde, formaldehyde, aceto acetic ester, B-oxybutyric

acid, creatinin, lactic acid, formic acid and inulin reduce in acid but not

in alkaline mixtures of sodium selenite. Methyl alcohol and ethyl alcohol

reduce sodium selenite strongly acidified with sulfuric or with hydro-

LEVINE, BIOCHEMICAL STUDIES OF SELENIUM 397

chloric. Ovxalic, citric, tartaric, malic and salicylic acids, benzaldehyde,

cinnamic aldehyde, and salicyl aldehyde reduce neither in acid nor in

alkaline mixtures.

The results show that monosaccharides readily reduce an alkaline solu-

tion of sodium selenite. The pentoses give readier and more profuse

reduction than the hexoses and the reducing disaccharides. Of the

pentoses, xylose yields most profuse reduction. Among the hexoses,

levulose and galactose reduce more readily than dextrose, and galactose

less readily than levulose. Among the disaccharides only those having a

free carbonyl group reduce. Maltose and lactose show reduction, but

sucrose does not. Raffinose, cellulose, starch, dextrin, glycogen, inulin

also do not reduce.

In order to test the influence of acidity or alkalinity upon the re-

duction, of sodium selenite, nineteen reagents were made up. One

consisted of sodium selenite neutralized with sulfuric acid. Another

consisted of sodium selenite, which reacts alkaline. ‘To nine other sele-

nite solutions were added sodium hydroxid, potassium hydroxid and Ro-

chelle salts, sodium bicarbonate, sodium carbonate, and sodium citrate,

sodium tetraborate, sodium silicate, disodium hydrogen phosphate. Eight

reagents were acidified by the addition of one of the following: potassium

bisulfate, sodium dihydrogen phosphate; hydrochloric, nitric, sulfuric,

phosphoric, citric, or tartaric acid. When these reagents were heated

none reduced, even on complete evaporation, except the one containing

citric acid and the one containing tartaric acid. These two reagents also

deteriorated after standing several months. Experiments with these

reagents were carried on at 37.5° C. and at 100° C. Solutions (0.5

per cent) of arabinose, rhamnose, xylose, glucose, fructose, galactose, su-

crose, maltose, lactose, glycogen, starch, dextrin,. inulin, raffinose, mucic

acid, lactic acid, formic acid, acetone and formaldehyde were used.

Three cubic centimeters of the solution to be tested were mixed with

two cubic centimeters of the selenite reagent and toluol added. The

tubes were incubated at 37.5° C., and examined from time to time.

Controls were run with the Fehling and the Fehling-Benedict reagents.

The reagents containing sodium hydroxid and potassium hydroxid

(selenite and Fehling) were the first to show reduction at 37.5° C. The

Fehling reagent reduced more quickly than the Fehling-Benedict. Gly-

cogen, starch, dextrin, inulin and raffinose reduced acidified solutions of

sodium selenite only at the end of four days. Alkaline solutions were not

affected. Formic acid, lactic acid, formaldehyde reduced in acid solutions

only. Acetone profusely reduced acid solutions, and very faintly reduced

alkaline solutions. The reagent acid with nitric showed no reduction,

388 ANNALS NEW YORK ACADEMY OF SCIENCES

except in the case of acetone. Neutralized sodium selenite proves to be

a very ineffective indicator for reduction. The presence of sodium teira-

borate inhibits to a very striking extent the reduction of sodium selenite.

The reagent containing 2% sodium selenite, 10% sodium citrate and

10% sodium carbonate has been tested with reducing sugars. Reduction

with this reagent takes place, at 100° C., in one minute or even less. At

first a deep chlorine-yellow color is developed. After standing a minute

or two this color gives way to a light wine-red tint, then to a dense brick-

red precipitate, which fills the volume of the tube. A 0.02% solution of

glucose yields fair reduction, and in a 0.01% solution the reaction is still

evident, but faint. Solutions to be tested must be alkaline, and must not

contain potassium cyanid or oxidizing agents (free halogen, hydrogen

peroxid, potassium permanganate, potassium bichromate). Sugar-free

urine gives a positive reaction when it is acidified with hydrochloric. This

positive reaction is probably due to acetone substances and creatinin,

which decidedly reduce acidified solutions of sodium selenite. Proteins,

uric acid and creatinin do not interfere with the alkaline sodium selenite

reagent.

Minute amounts of selenium in the form of selenite ion can be de-

tected by a procedure similar to that of the Marsh test for arsenic. One

milligram of selenium dioxid yields a characteristic dull red selenium

mirror, soluble in oxidizing agents.

REDUCTION OF SELENIUM COMPOUNDS IN THE LIvING ORGANISM

PLANT SUBSTANCES

Plant substances (apple pulp and potato pulp) bring about the reduc-

tion of sodium selenite to brick-red selenium. An alkaline reaction

favors the reduction process. Boiled material induces no reduction or

very faint reduction.

YEAST

Yeast reduces selenious acid, selenic acid and sodium selenite. The

red selenium deposits in the cell-body so that only the cells themselves

are pigmented, while the liquid above remains colorless. The cells can

be decolorized by washing with potassium cyanid.

UNBOILED MILK

Unboiled milk in contact with a few drops of sodium selenite (room

temperature) will show reduction within one to three days; boiled milk

will show no evidences of reduction. Unboiled milk protected with

toluol does not reduce as readily as milk exposed to the air.

LEVINE, BIOCHEMICAL STUDIES OF SELENIUM 389

ANIMAL TISSUES

Animal tissues also reduce sodium selenite. Fresh liver, spleen, heart,

lung, kidney, pancreas, small intestine, large intestine and stomach, in-

cubated in the presence of toluol at 37.5° reduce a 0.5% solution of

sodium selenite within twenty-four hours. The liver and spleen reduce

very quickly and most profusely. Portions of striped muscle, testicles,

thyroid, submaxillary gland and lingual glands give negative results.

Toluol-preserved, unfiltered extracts, first kept at room temperature for

twenty-four to thirty-six hours and then treated with sodium selenite

showed reduction within twenty-four hours, only in the case of liver,

yancreas and small intestine. This result indicates that the reducing

power of the tissues disappears in the course of time. That the reduc-

tion is due to cell activity or enzyme activity is evident from the fact

that tissues after being heated on a water-bath for about ten minutes

fail to yield reduction. Fresh liver, however, heated or unheated, re-

duces sodium selenite. ‘This reaction is due to the presence of chemically

reducing substances such as carbohydrates. Tissues to which an excess

of chloroform has been added show no reducing power. Chloroform itself

does not inhibit the reduction of sodium selenite by pure sugar solutions.

Filtered chloroform extracts also do not reduce, although the liver extract

was found to reduce on the first or second day, and the spleen on the

third or fourth day. The other extracts failed to show reduction, even

after two weeks. Reduction is probably due to an insoluble enzyme closely

connected with the living cell, that is, an endo-enzyme. Selenious acid

and selenic acid are also reduced, while sodium selenate is not reduced.

SELENIUM COMPOUNDS INJECTED INTO THE ANIMAL SYSTEM

Selenium compounds, injected into the animal system, undergo re-

duction. Some of the selenium escapes from the organism in the form

of a volatile organic selenid, some is precipitated, mostly extracellularly,

in the tissues as dark red-brown granules. ‘he liver and spleen contain

by far the largest amount of deposited selenium. ‘The microscopic ex-

amination of the histologically stained tissues of a dog that had received

2 mg. selenium dioxid per kilo, of body weight, revealed the presence of

selenium in these two organs. With sodium selenite, selenium was found

widely distributed in the liver, spleen, kidney, lung, pancreas, heart,

stomach and intestine. These tissues showed marked reductions within

the regions of hemorrhagic clots. Other tissues showed reduction only

within such areas. A lethal dose of selenic acid brought about deposition

of the granules in the spleen and liver. The lungs, which were found

to be extremely congested, also showed reduction. Selenium pigmenta-

390 ANNALS NEW YORK ACADEMY OF SCIENCES

tion with potassium selenate was slight in comparison with that produced

with sodium selenite and but few granules were found scattered in the

spleen, pancreas and liver. In the case of potassium selenocyanate, re-

duced selenium was found in the liver, spleen, lung, kidney, pancreas,

heart, brain (only in hemorrhagic spots), muscle (only im similar spots)

and stomach.

The reduction of selenium compounds seems to be a detoxicating

process, since selenium itself is hardly poisonous. It seems evident that

while sulfur compounds, such as sodium sulfite, are oxidized in the body

to sulfate, the corresponding selenium compounds, such as sodium sele-

nite, suffer reduction.

LIVING BACTERIA

Living bacteria bring about the reduction of selenious acid, selenic

acid and sodium selenite. The precipitated selenium follows the path

of bacterial growth. Granules of selenium can be observed under the

microscope within the bacterial cell. The medium used should not con-

tain reducing chemical substances, such as glucose or lactose. Sodium

selenaie is not reduced. Due to decomposition induced by acids, selenium

may be also deposited from potassium selenocyanate. This happens to

be the case with B. colt.

Reduction is proportional to the intensity of growth. Selenious acid,

selenic acid and, less markedly, sodium selenite do not favor growth.

The amount of retardation depends upon the nature of the organism.

Streptococcus pyogenes was found to be more sensitive than Bacillus colt.

The bacilli of symptomatic anthrax, edema, and tetanus are retarded very

markedly in growth. Sodium selenate and potassium selenocyanate do

not retard growth.

Selenium dioxid or sodium selenite cannot be used as a differential

test between xrobic and anzrobic organisms, since the latter class also

bring about reduction. There seems to be no specific relationship be-

tween selenium reduction and hydrogen sulfid production as Gloger

maintained, since micro-organisms, such as B. acidi lactict, B. dipth-

therve, B. pseudo-diphtherie, B. tuberculosis that produce no hydrogen

sulfid or only traces, were capable of reducing selenium dioxid or sodium

selenite.

For practical purposes selenium dioxid or sodium selenite in concen-

trations of 1:50,000 or 1:25,000 can be used to demonstrate bacterial

reduction. This phenomenon can also serve as a qualitative test for the

selenite ion, even if the selenious acid or the sodium selenite is present

in culture media in such small concentrations as 1:100,000.

LEVINE, BIOCHEMICAL STUDIES OF SELENIUM 391

EFFECT OF SELENIUM COMPOUNDS UPON CATALASE AND OTHER

ENZYMES

CATALASE

The compounds of selenium employed in enzyme investigations were

selenium dioxid (selenious acid), selenic acid, sodium selenite, sodium

selenate and potassium selenocyanate. The effect upon catalase was

determined as follows: Healthy, normal dogs were bled to death from

the femoral artery, using cocaine as a local anesthetic. Weighed

amounts of defibrinated blood and tissues were ground with sand and

mortar, treated with 40 c. c. distilled water and 10 c. c. chloroform and

permitted to extract twenty-four hours. ‘These served as controls. Equal

amounts of blood and tissue were treated in the same manner, except that

the distilled water was substituted by a 1/20% solution of selenium acid

or a 1/10% solution of a selenium salt. After the extraction period a

definite volume of the filtrate was treated with 5 c. c. Oakland dioxygen

and the catalytic powers of the filtrate as measured by the volume of

oxygen evolved, determined for every thirty seconds. The liver, kidney

and blood showed the greatest catalytic activity.

The compounds of selenium investigated had a marked inhibitory

effect on catalytic activity. Thus ten grams defibrinated blood evolved

52 ¢. c. oxygen in 7 minutes, another sample of the same blood treated

with 0.05% selenium dioxid evolved 49.8 c. c. in 21 minutes, a third

sample of the blood treated with 0.05% selenic acid evolved 52.6 c. c. in

14 minutes.

Blood from another dog yielded 44.8 c. c. in 9 minutes, another sample

of the same blood treated with 0.1% potassium selenocyanate evolved

37.7 c. c. In 22 minutes. Control liver extract yielded 52.7 c. c. in 2

minutes ; selenited liver extract yielded 50.2 c. c. in 8.5 minutes; selenated

liver extract, 49 c. c. in 4 minutes. Another control liver gave 45.5 ec. ¢.

in 5.5 minutes, while another sample of the liver treated with potassium

selenocyanate yielded 41.7 c. c. in 13.5 minutes. Generally speaking,

the blood, liver, kidney, lung and spleen showed marked decrease in

catalytic activity, the decrease sometimes being as much as 60% or over.

Compared on the basis of equipercentages, it was found that selenium

dioxid was more harmful than selenic acid, and that sodium selenite

produced greater inhibition than sodium selenate. It is interesting to

note, however, that colloidal selenium (prepared by the reduction of

sodium selenite by glucose) brought about a slight acceleration in catalase

activity.

392 ANNALS NEW YORK ACADEMY OF SCIENCES

Tissues of dogs killed with selenious acid, with selenic acid or with

potassium selenocyanate showed no reduction in catalase values. This

fact points to the decomposition of the selenium compounds injected,

with the formation of substances that had no inhibiting effect on catalase

action.

SALIVARY AMYLASE

The influence on salivary amylase was determined by Wohlgemuth’s

method. Small amounts of sodium selenite (neutralized) and sodium

selenate (0.05% to 0.1%) had a slight effect on ptyalin. In the presence

of 0.05% sodium selenite and, more markedly, in the presence of 0.05%

sodium selenate the activity of the amylase seemed slightly increased.

The presence of potassium selenocyanate in the saliva or in the urine

interferes with the Fehling-Benedict reduction test.

PEPSIN

The results obtained with pepsin showed that selenious or selenic acid

could replace the hydrochloric acid of the gastric juice. Selenious acid

was slightly inhibitive, while selenic acid resembled sulfuric acid in the

marked inhibition which it shows towards proteolysis. Sodium selenite

(neutralized) and sodium selenate (0.01% to 0.2%) had little or no

effect on peptic activity. Higher concentrations produced inhibition,

this being more marked with sodium selenite than with sodium selenate.

Potassium selenocyanate, even in minute amounts, inhibits digestion. this

probably being due, in part, to the mechanical interference of the brick-

red precipitated selenium, which completely covered the fibrin, or pos-

sibly to the presence of the compound that results from the acid decom-

position of potassium selenocyanate.

TRYPSIN

Slight amounts of sodium selenate and potassium selenocyanate had

no effect on tryptic activity. Neutralized sodium selenite inhibited even

in small quantities.

RENNIN

Sodium selenate and potassium selenocyanate (0.05%-0.5%) had no

influence on rennin. Coagulation was retarded by concentrations of

neutralized sodium selenite above 2%. Sodium selenite and sodium

selenate had but-a-slight effect on the souring of milk. Potassium seleno-

cyanate showed an inhibitory effect, the amount of inhibition being

directly proportional to the concentration of the salt.

LEVINE, BIOCHEMICAL STUDIES OF SELENIUM 393

PANCREATIC LIPASE

Sodium selenate and potassium selenocyanate had no effect on pan-

creatic lipase. Sodium selenite had a shght inhibitory effect, which was

proportional to the concentration. Experiments carried on with selenium

dioxid and with selenic acid showed a marked increase in acid content.

The high results can be attributed to the hydrolytic stimulation of these

selenium acids.

ALCOHOLIC FERMENTATION

Selenium dioxid in concentrations of 0.5% had marked inhibitory ac-

tion on alcoholic fermentation. In a 2.5% solution fermentation occurred,

but was suppressed entirely in a 3% solution. Sodium selenite (alkaline)

inhibited the evolution of carbon dioxid. Selenic acid, even in a con-

centration of 0.04% exercised an inhibitory influence over zymase. Very

little carbon dioxid was evolved with a 0.2% solution, while solutions

containing 0.5% or over gave no evidence of carbon dioxid formation.

Sodium selenate accelerated alcoholic fermentation. The effect produced

by potassium selenocyanate seemed to be variable, although, in general,

the inhibition produced by concentrations lower than 2.5% was slight.

Toxic EFFECTS oF SELENIUM COMPOUNDS

Selenium compounds are toxic to both plants and animals. Even con-

centrations of 0.01% inhibit germination and growth. Sodium selenate

is the least toxic of the compounds investigated. Beginning with the

most poisonous the sequence of toxicity for animals is hydrogen selenid,

selenious acid, selenic acid, sodium hydrogen selenite, sodium selenite,

potassium selenocyanate, sodium selenate and free selenium. Potassium

selenocyanate is more toxic than the corresponding potassium sulfo-

cyanate.

In very minute doses selenium compounds have no effect on blood

pressure. In comparatively larger doses a marked fall in blood pressure

is observed in the case of selenious acid, selenic acid, normal sodium

selenite, sodium hydrogen selenite and sodium selenate, although with

potassium selenocyanate a considerable rise in blood pressure is induced.

Selenium compounds, even in small doses, have a marked effect on

respiration. Breathing becomes prolonged, deep and labored, and as the

dose is increased respiratory paralysis sets in even before the heart stops.

Pulmonary edema, accompanied by exudation of large volumes of yellow-

ish fluid, precedes death in the case of selenious acid, sodium hydrogen

selenite, sodium selenite and selenic acid.

394 ANNALS NEW YORK ACADEMY OF SCIENCES

The characteristic odor of a volatile selenium compound (methyl

selenid ?) is scented in the animal’s breath almost immediately after

intravenous injection and about ten or fifteen minutes after subcutaneous

injection. In one case, where the dose was five times the lethal dose,

the odor was not evident. Another characteristic effect of selenium

poisoning is the withdrawal of free hydrochloric acid from the stomach.

To account for the toxicity of the selenates we put forth the proposition

that the selenates are reduced to selenite and finally to free selenium.

To explain the toxic properties of sodium arsenate, Binz and Schulz

claim that the arsenate is reduced in the animal body to the more toxic

arsenite.

The selenium compounds investigated do not precipitate proteins from

their solutions.

The author gratefully acknowledges his indebtedness to Professor

William J. Gies for kind assistance, valuable suggestions and helpful

criticism.

Biochemical Laboratory of Columbia University,

College of Physicians and Surgeons.

PLATE V

ILLUSTRATIONS OF BIOLOGICAL REDUCTIONS OF SELENIUM COMPOUNDS

Fic. 1.—B. coli grown on a sodium selenite culture medium. The colonies

appear as red spots.

Fic. 2.—Another culture of B. coli grown on a slant.

. 3.—Section of the liver of a dog that had been subcutaneously treated

with 2 mg. selenium dioxid per kilo of body weight. - The chocolate

red granules represent deposited selenium.

. 4— Stab culture of streptococcus pyogenes.

a4 ANWALS NOW YORK ACADBMY GE AGIENGES — :

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tae hat the arsenate is 5 redueed int i the animal ‘ot

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