OFFICERS, 1911

President—Franz Boas, “olumbia University |

Vice-Presidents—Gronrcr F. Kunz, Frepertc A. Lucas, .

R. S. WoopwortH, WILLIAM CAMPBELL

_—- Recording Secretary—EpMunD Otis Hovey, American Museum —

seo Corresponding Secretary—Henry HE. CRAMPTON, American ‘SMoeum ‘

——s- Preasurer-—EMERSON McMituin, 40 Wall Street fie et .

ss Librarian Rate W. Towne, American Museum a

ANA SECTION OF GEOLOGY AND MINERALOGY

Chairman—GeorceE F. Kunz, 401 Fifth Avenue

Secretary—Cuar.zs P. Berkey, Columbia University

“ees _ SECTION OF BIOLOGY

— - Chatrman—Freveric A. Lucas, American Museum.

» Secretary—L. Houssakor, American Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—WiLt1am Campsett, Columbia University

Secretary—Epwarp J. THATCHER, Teachers College

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. S. WoopwortH, Columbia University

Secretary—F RepERIc LyMANn WELLS, Columbia University

eet The sessions of the Academy are held on Monday evening at 8: 18 |

oa o’clock from October to. May, inclusive, at the American Museum ;

Natural History, 77th Street and Central Park, West. |

[Annats N. Y. Acap. Scr, Vol. XXII, pp. 1-8, Pl. I. 3 April, 1912.]

ON SOME INVERTEBRATE FOSSILS FROM THE LYKINS

FORMATION OF EASTERN COLORADO?

By Grorce H. Girty

(Read by title before the Academy, 5 February, 1912)

The fossils which form the subject of the following account were col-

lected by Mr. Roy M. Butters and kindly placed by him in my hands for

study. They were obtained in the Lykins formation of Colorado and

represent a horizon in the Paleozoic higher than any at which fossils have

heretofore been found along the eastern flank of the Front Range.

A detailed account of the stratigraphic relations and correlation of the

Lykins formation has been prepared and will shortly be published by Mr.

Butters. To a manuscript of this report, which I have been permitted to

read, I largely owe the following data which seemed essential to the

understanding of this limited but interesting fauna.

The “Red Beds” of the Front Range in Colorado have been variously

classified and named. Their nomenclature and synonymy is, therefore,

rather complicated, but as a general statement, it may be said that the

Wyoming formation of Emmons has been divided into three formations,

of which the Lykins is the highest. Below the Lykins, there occurs a

series of strata (the lower Wyoming) which are now known as the Foun-

tain and Lyons formations, while above the Lykins is the Morrison for-

mation. The Lykins, therefore, belongs in the upper “Red Beds” of this

area. The Fountain has furnished more or less conclusive paleontologic

evidence of Pennsylvanian, or at least of upper Carboniferous, age, while

the Morrison has long been known to be Mesozoic. The Lykins forma-

tion, from which fossils have not hitherto been known, has usually been

assigned to the Triassic, but the evidence herewith presented seems to

show conclusively that the formation, or at all events that portion of it

from which the fossils were obtained, is Paleozoic. Provisionally, I am

assigning the Lykins fauna to the Permian, though more on account of

its position at the top of the Paleozoic section than on account of any

very close resemblance either to the Permian of Russia, the more or less

doubtful Permian of our Western States or the Permian as distinguished

from the Pennsylvanian of the Mississippi Valley. The only fauna in

2 Published by permission of the Director of the U. S. Geological Survey.

9 ANNALS NEW YORK ACADEMY OF SCIENCES

Colorado which conspicuously resembles the Lykins occurs in the Rice

formation of the San Juan region, and the Lykins and Rico are tenta-

tively placed in correlation, in spite of the fact that many of the Rico

forms are as yet not known in the Lykins and some of the characteristic

Lykins forms are not known in the Rico.

The character of the Lykins formation, as would naturally be supposed,

changes from point to point. A well-exposed section at Heygood Canyon

which is fairly representative consists, according to Mr. Butters, of sand-

stone and shale with some beds of sandy limestone. The sandstones are

pink or red and mostly soft, while the shales are red. The thickness of

the Lykins at this point is 816 feet.

A little south of Heygood Canyon on the north slope of Table Moun- _

tain were made three collections of fossils (lots 3264, 3265 and 3266).

They occur about 300 feet above the base of the formation and about 25

feet above a 25-foot bed of gypsum. They contain the same species, viz:

Myalina perattenuata, Myalina wyomingensis, Alula squamulifera and

Murchisonia buttersi. The matrix is a compound of fine sand and clay

with more or less lime, the color being a rather light brownish gray.

Another collection was made near Stout, Colorado, from a red calcare-

ous sandstone (lot 3262). Only two species are present, Myalina wyo-

mingensis and M. perattenuata. The horizon is about 30 feet above the

“crossbedded sandstone” * in the basal member of a local group of cal-

careous strata 50 to 60 feet thick consisting of thin limestones, shales

and sandstones.

The last collection (lot 3263) was made at Perry Park, one-fourth of

a mile south of the lake, in a band about 6 inches thick near the base of

the “crinkled sandstone.” * The rock is whitish in color and very fine in

texture, apparently a mixture of lime and clay and sand. In this collec-

tion, I identify Myalina wyomingensis, Myalina perattenuata, Alula squa-

mulifera, Alula gilberti? and Pleurophorus sp. —

Lot 3263 occurs at a higher horizon than 3262 and recalls, especially

by the peculiar and characteristic species Alula squamulifera, the fauna

of the northern group of collections (lots 3264, 3265, 3266), from which,

however, it is separated geographically by atong distance.

The fauna of the Lykins formation is, so far as known, very limited,

consisting of only six species, and in addition to describing the two spe-

cies, which are new, it has seemed desirable to remark briefly upon the

other forms.

* Colorado Geol. Survey, First Report, pp. 168-9. 1909.

3U. 8. Geol. Survey, Bull. 265, p. 25. 1905.

GIRTY, INVERTEBRATE FOSSILS FROM COLORADO 3

Alula gen. nov.

Shell soleniform, very transverse. Beak strongly anterior, but not terminal.

Upper and lower margins contracting posteriorly. Posterior outline obliquely

truncated. Umbonal ridge angular, with a second plication on the post-car-

dinal slope. Surface elegantly sculptured by fine radial costz surmounted by

fine, closely arranged scales or interrupted concentric lamelle. Post-cardinal

slope without radial costs, but with similar squamose ornamentation. On the

interior, the right valve has a single long, plate-like anterior tooth, a posterior

tooth of similar character, with possibly a small rounded tooth at the umbo.

Corresponding structures appear to be developed in the left valve. A large

anterior scar is indicated.

Type, Alula squamulifera.

In a general way, these shells suggest a very transverse type of Paral-

lelodon, and I believe that they belong to the same family, though clearly

representing a distinct genus. In configuration, they differ from Paral-

lelodon in contracting posteriorly instead of anteriorly ; in not having the

angular and projecting anterior extremity, and in possessing a second

plication between the umbonal ridge and the cardinal border. Interiorly,

they differ in having a single posterior tooth; in having a linear anterior

tooth similar to the posterior one, and probably in lacking the flexuous

dental arrangement at the umbones. A certain resemblance to some spe-

cies of Plewrophorus exists in the angular umbonal ridge and the post-

umbonal fold, but the Plewrophori are not radially striated, and, while

they have a similar posterior tooth, the remainder of the dentition is

quite different.

It is not certain that any American species other than the type can be

referred to this genus, but, if so, they are probably to be found among

the forms which I provisionally included under Pleuwrophorella. A re-

semblance to Alula squamulifera, more or less marked, is found in P.

gewmta, P. gilberti, P. lanceolata and Allerisma (Pleurophorella?) re-

flecum. Of these, the most similar is P. gilberti. Typical Pleurophorella,

as exemplified by P. papillosa, is probably safely distinct, although its

internal characters are as yet unknown, because of the deeply introverted

lunule and the escutcheon, both characters apparently wanting to Alula,

and because of the absence of radiating costz in the sculpture, although a

somewhat similar feature exists in the characteristic papille, which show

a tendency to radial arrangement. There is, however, scarcely any com-

parison in this item of sculpture. As a provisional arrangement, I am

removing to the present genus A. gilberti, A. geinitzi and A.? lanceolata.

Allerisma reflecum, in spite of a general resemblance to this series of

forms, probably has quite different, although indeterminate, relations,

distinctly not with typical Allerisma. As a result of a better knowledge

4 ANNALS NEW YORK ACADEMY OF SCIENCES

of Allerisma costatum and a renewed consideration of its characters, I

believe that my original estimate of its relationship to Pleurophorella

papillosa was erroneous. The strong concentric plications which stop

abruptly at the umbonal ridge indicate a different type of shell. It is

somewhat doubtful whether a papillose surface is a real character of A.

costata, which is apparently a much flatter shell, with thin test and pos-

sibly different structure in the lunule and escutcheon. It clearly does not

belong with Alula, however, but has all the superficial characters of

typical Sanguinolites.

Alula squamulifera

Shell rather small, very transverse. Width about 3.5 times the greatest

height. Greatest height near the anterior end at the umbo, which is situated

about one-sixth of the entire width back from the anterior margin. Ventral

border gently .convex in the anterior half, nearly straight or faintly concave

posteriorly. Dorsal outline gently concave or nearly straight, contracting pos-

teriorly with the ventral. Posterior outline oblique and more or less sharply

truncate. Anterior outline straight above, strongly rounding below. Convexity

usually rather high, though variable, sometimes rather tumid in the umbonal

region. Beaks large, prominent and incurved, situated relatively close to the

anterior extremity. Umbonal ridge prominent, usually strongly angular toward

the posterior end, more obscure in the umbonal region. The post-cardinal

slope is divided by a second plication about intermediate between the umbonal

ridge and the cardinal line, above which the narrow strip of shell is nearly

horizontal. Surface marked by fine, radiating ribs which are confined fe the

portion of the shell below and in front of the umbonal ridge. This Svu.pr.ure

might better be described as made by narrow stris, the elevations between

which are covered with closely arranged, fine, flat scales, which recur at equal

intervals on adjacent ribs and have also the appearance of interrupted con-

centric lamelle. The ribs are more than radiating rows of scales, since the

spaces between them are depressed. The scales are sometimes more or less

eurved with the convex side uppermost, especially at the anterior end, where

they are replaced by two or more rows (the radiating arrangement often not

being apparent) of minute spines or papille. Apparently, these spines become

more or less compressed toward the middle of the shell and then coalesce at

their edges. If they are not quite in alignment, the curved appearance noted

above results. The post-cardinal slope, which, as already mentioned, lacks

radiating ribs, is nevertheless marked by these flattened scales, which tend to

be arranged in concentric rows without, however, becoming connected into

continuous lamelle. No radial arrangement is here apparent.

The internal structures are imperfectly known. The right valve bears two

linear teeth, one before and one behind the beaks. The posterior tooth is long,

about two-thirds the entire length back of the beaks. The anterior tooth is

much shorter, about one-half the length of the anterior outline. Whether a

small cardinal tooth was developed between these at the umbo is not clearly

shown, but such a structure is indicated. In the left valve, there appear to be

linear sockets corresponding to the teeth of the right. A large anterior scar is

indicated,

GIRTY, INVERTEBRATE FOSSILS FROM COLORADO 5

Of described species, this appears most closely to resemble A. gilberti,

though it is not certain that the two are congeneric. The chief difference

of a possible generic character lies in the fact that White’s figure ap-

pears to represent A. gilbertt as having a well-marked escutcheon, a

structure probably not present in A. squamulifera. Specifically, the

latter appears to be a more slender form, more convex, and with a sharper

umbonal ridge (these characters, however, may be enhanced by compres-

sion in the Colorado form). It is also distinctly, though finely, costate,

although A. gilberti in fact is covered with granules arranged in rows, so

as to resemble minute radiating lire.

Horizon AND Locality: Lykins formation; Heygood Canyon (lots

3264, 3265, 3266) and Perry Park (lot 3263), Colorado.

Alula gilberti White?

Alula squamulifera is abundant in lot 3263, but specimens are in an

unsatisfactory condition of preservation. Many of them show a lower

convexity and less angular umbonal ridge than the types. One example

is sufficiently shallow, broad and ill-defined as to umbonal ridge to re-

semble Allerisma gilbert rather closely. The sculpture is obscure but

presents suggestions of radiating coste or of rows of papille. The de-

pressed specimens which are provisionally placed with A. squamulifera

appear to show a gradation toward but not into the only one referred to

White’s species, and the facts which I have been able to observe leave me

in doubt as to whether we have three species of not necessarily generically

identical shells, or a fairly continuous series of mutations with A. squa-

mulifera at one end and A. gilberti (or the form here identified as such)

at the other.

Horizon AND LocaLity: Lykins formation; Perry Park, Colorado

(lot 3263).

Myalina wyomingensis Lea

Myalinas are extremely abundant in four of the five collections exam-

ined, but most of the specimens are small. They vary in specific charac-

ter. Some of the larger and more typical specimens agree in every

determinable character with forms from the Rico formation of the San

Juan region which I identified as Myalina wyomingensis.t. The great

majority are of much smaller size, more like the form from Ouray which

I somewhat provisionally called MW. cuneiformis.2 They naturally have

the anterior lobe less strongly developed than the larger or mature exam-

ples which accompany them. They seem as a rule to be less strongly

4U. S. Geol. Survey, Prof. Paper 16, Plate VIII, Fig. 8. 1903.

5 Ibid., Plate VIII, Figs. 16 and 17.

6 ANNALS NEW YORK ACADEMY OF SCIENCES

oblique than the type specimens of M. cuneiformis, though some of them

have the lobe scarcely more apparent. I am regarding part of these small

specimens as being young examples of M. wyomingensis, and this may

also be the true relationship of the Ouray specimens referred to cunet-

formis. Typical cunetformis should probably be kept distinct for the

time being.

Horizon AND LocaLity: Lykins formation; Heygood Canyon (lots

3264, 3265 and 3266), Stout (lot 3262) and Perry Park (lot 3263),

Colorado. :

Myalina perattenuata Meek and Hayden

The Myalinas of the Lykins formation in addition to showing varia-

tion in the amplitude of the posterior wing vary conspicuously in the

development of the anterior lobe. Some specimens have scarcely any

perceptible development of this feature. These, although they are not

sharply distinguished from the typical J/. wyomingensis, | am separat-

ing as a different species under the title M. perattenuata. A similar phe-

nomenon was observed in the Myalinas of the Rico formation of the San

Juan region, and a similar course was pursued in regard to them. These

Lykins specimens, however, are for the most part much smaller than

those from the Rico formation and in this character approximate M.

cuneiformis, but most of them are distinctly less oblique. A not very

considerable breakage along the hinge line of these small shells however,

or a concealment of the true outline in that region, makes an appreciable

difference in their apparent obliquity.

Horizon AND Locality: Lykins formation; Heygood Canyon (lots

3264, 3265 and 3266), Stout (lot 3262) and Perry Park (lot 3263),

Colorado.

Pleurophorus sp.

A very imperfect internal mold showing best the impression of the

hinge structures in the umbonal region, where they possess the charac-

teristic dental arrangement of Plewrophorus. For the rest, there is indi-

cated a transverse, oblong shell of medium size with rather strongly

projecting anterior end.

Horizon AND LocALiITy: Lykins formation; Perry Park, Colorado

(lot 3263).

Murchisonia buttersi sp. nov.

Shell of medium size, slender, with high, many-whorled spire. Length of the

type specimen as restored about 25 mm. Diameter of final whorl 11 mm.

Number of volutions 10. Volutions angular with a thick, prominent carina

situated considerably below the middle, the height of the upper zone being to

that of the lower about as 2 to 1. Upper and lower zones more or less planate

GIRTY, INVERTEBRATE FOSSILS FROM COLORADO 7

and standing at approximately a right angle to one another. The lower is

gently concave, more so than the upper, although the upper spoons outward as

it approaches the carina. Suture deeply depressed.

The most conspicuous superficial feature consists of narrow angular cost,

leaving between them broad shallow interspaces, which cross the upper portion

of the yolutions transversely or in a direction longitudinal to the shell as a

whole. They are straight, but are slightly oblique, retrally directed from

above downward. These plications are’ perhaps restricted to the three or four

older volutions, and there is some irregularity in their arrangement. They

die down before reaching the carina. In addition, the whole surface of the

upper zone is marked by microscopic transverse and revolving lire producing a

more or less cancellated effect. The revolving lire are rounded, closely

arranged and prone to be wavy. The transverse lirze are finer, sharper and

more irregular, more of the nature of incremental lamelle, and the coste may

perhaps be looked on as fascicles of these markings. The lower zone of the

volution is marked similarly to the upper, but the angular cost are less strong.

They have a slight forward obliquity from the carina. There appear to be two,

possibly more, strong rounded revolving lire on the final volution at a point,

as it would appear, about half-way down from the carina, and the volutions so

embrace as to leave about two of these lire visible above the deeply sunk

suture. The final volution is not well shown by the specimens examined, so

that the sculpture below these two lire, the relative distance at which they

occur below the carina, the shape of the aperture, ete., are not known. The

carina is the site of the slit band. The band is occupied by two rather coarse,

rounded lirz, separated by a narrow stria and appears to be defined by two

delicate lamellose lines, one above and one below, bounded on the median side

by slight striz. The two revolving lire which occupy the whole of the band

and are more projecting than the edges are rendered nodose by the cost#e

described as crossing the upper and lower surfaces of the volution. That is,

the swellings occur where the cost would cross them, but the coste are evan-

escent on the upper surface near the band, and the nodes are much more

prominent than the costze and much more elongated spirally.

Tn its specific relations, this shell is most nearly related to Murchisonia

lasallensis and M. terebra. It differs from both in the presence of trans-

verse plications. From terebra, which seems to be more nearly related

than the other, it apparently differs also in having the carina containing

two crenulated lire instead of one, in having two revolving lire just

above the suture and in other details of sculpture.

Generically, this shell can hardly be classed with typical Murchisoma,

though it belongs to a group frequently cited under that genus. In some

important respects, it is comparable with such representatives of the

genus Worthenia as W. tabulata. This is especially true of the structure

of the slit band, which seems to be identical in both. Given a much

higher spire and more gradually enlarging volutions, with some modifica-

tions in the modeling of the whorls, especially the lower part, it is easy

8 ANNALS NEW YORK ACADEMY OF SCIENCES

to conceive how such a configuration as that of M. butters: might be

evolved from that of Worthenia tabulata. The sculpture also appears to

be of the same general character, the most essential difference being the

development of transverse costz and of revolving lire more prominent

than the rest on the lower half of the inferior zone. Some important

characters of M. butters: are still unknown, but if these show no addi-

tional differences, it may prove to be a rather extreme form of Worthenia.

Horizon AND LocALiry: Lykins formation; Heygood Canyon, Colo-

rado (lots 3264 and 3266).

PLATE I

LYKINS FOSSILS

Myalina perattenuata (p. 6)

Fig. 1. A large right valve referred to this species.

2. A left valve of more nearly the average size.

Lykins formation, Heygood Canyon, Colorado (lot 3266).

Myalina wyonrvingensis (p. 5)

8. A large left valve.

Lykins formation, Heygood Canyon, Colorado (lot 3265).

Alula squamulifera (p. 4)

4, Side view of an internal mold of a right valve.

4a. A view obliquely down on the cardinal margin of the same specimen,

showing the impression left by the linear anterior and posterior

teeth, x 2.

5. Squeeze of a right valve showing the surface characters.

5a. Same, x 2. Hyen with this magnification, the fine squamose character

of the costz cannot be shown.

Lykins formation, Heygood Canyon, Colorado (lot 3266).

Alula gilberti? (p. 5)

6. Side view of a doubtfully identified right valve.

Lykins formation, Perry Park, Colorado (lot 3263).

Murchisonia buttersi (p. 6)

7. Side view of a squeeze made from the type specimen.

8. Another squeeze made from the same specimen, x 2.

Lykins formation, Heygood Canyon, Colorado (lot 3264).

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ANNALS N.Y. ACAD. Sci. VOLUME XXII, Plate |

FOSSILS FROM THE LYKINS FORMATION

nes ACADEMY OF SCIENCES

”

“*

‘(bron OF Naruran History, 1817-1876) —

ublications of the Academy consist of two series, viz.:

r NS Ses

oe is led that — shall be devoted to

icone are sent fre ¢ to Fellows and Active Members. The .

s are sent to Honorary and Corresponding Members desiring them. a

Lees oe Inquiries concerning current and back numbers of

“Ge LIBRARIAN,

New York Academy of Sciences,

. eare of

American Museum of Natural History,

New York, N. Y. ae

a int

Joun D. HaseMan

i. NEW YORK Tee

PUBLISHED BY THE ACADEMY — es

31 May, 1912 :

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or NatruraL History, 1817-1876)

OFFICERS, 1912

President—Emrrson McMuttin, 40 Wall Street

Vice-Presidents—J. EDMUND WoopMAN, FREDERIC A. Lucas,

CHARLES LANE Poor, R. 8S. WooDwoRtH

Corresponding Secretary—Henry EH. Crampron, American Museum

Recording Secretary—Epmunp Otis Hovey, American Museum

-Treasurer—Hunry L. Douerty, 60 Wall Street

Librarian—RatpPu W. Tower, American Museum

EHditor—Epmunp Otis Hovey, American Museum

SHCTION OF GEOLOGY AND MINERALOGY

Chairman—J. E. WoopMan, N. Y. University

Secretary—CuHarLes P. Berkey Columbia University

SECTION OF BIOLOGY

Chairman—Freperio A. Lucas, American Musepm

Secretary—Wi.uiamM K. Grecory, American Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—CHARLES LANE Poor, Columbia University

Secretary—F. M. Prprrsen, College of the City of New York

SHCTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. 8. WoopwortH, Columbia University

Secretary—F RepEric Lyman WELLS, Columbia University

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 Rian

Natural History, 7th Street and Central Park, West

{Annas N. Y. Acap. Scr., Vol. XXII, pp. 9-112, pll. II-XVI. 31 May, 1912]

SOME FACTORS OF GEOGRAPHICAL DISTRIBUTION IN

SOUTH AMERICA

By Joun D. Haseman

| Presented in abstract before the Academy, 12 February, 1912|

CONTENTS

PaGcr

PLY POT SIGIR DE J ~ o'g.5 5 Sag SIs GS BnIene ono Gas STH One OOH IRE ec 10

Part I. Geology and topography of South America..................... 17

Disipuuion Of c<eolosical MHOTiZONS.: . 2.0.2 6s ee ce ee ee le eee ee 17

THEETGL IMMGSE So dest Ss Sooo Sete cae ence ee Cuca rer Ch eer tr ea tar Sena ean a 22

1S SRE VERISIGN 25

Eliane Alto and the Permian Inland Basin............2.0..5...06-- 27

JEASTE AIOE SSAA Sao 6 o Shs able ete OI SPO RSI a ee eae ea 30

Reversal of Rio Amazonas.......... BOE ae eRe dt says ee cach eyo Rls, ch eae 33

SHOENT [EMIPAGPY 6.5 5.5 Wh Soi AR res eTocs ee greege 38

Barriers to aquatic migration and conditions of environment......... 42

Part II. Distribution of the South American fishes and its bearing upon

alleged connections between South America and the eastern

VETNISTOLNERE. 5665 Sos ae eters ie BieceiaIcle Te Siar eae eae 50

1 CONST FES ETN OVA SCONE TN VES THIS] NSS ee 50

SL eneaNe te OMEN Ce La eee Me esg etn cf occa tle iis) atencacty S58 p16 isle whe, aoe hOacie ain 50

WISE CIE. CINAIECRERS iy ta Bis Sires Soe OCR a ee 55

Ne VOLO SAGA esol UN allem COUOMSi sere\s, sto. e8ewrs Mickc eoccs « aaa wsieie sues ols cre 55

Rio Uruguay and Rio Grande do Sul, including part of the lower La

LPVAVED! 5 5 5.9 A az ,4 d OOS BH ee Sg Ge EEN Re RE Cn ne eae re eee 62

Alta Parana and coastwise streams of eastern Brazil.................... 63

Sao Francisco and the Secca (dry) region of northeastern Brazil... ..... 64

the Earacsuay as part of the Amazonian complex................-2+0+5- 64

enue ETA DAC T AC OMI aye eye i oe ea css csece seecce beet biceus 66

ORAM whe SOUUM AMeLIGHM TSHES. . oo... clos ob dc cles cece dates i)

Summary of the most important data which have been used to support the

view that South America and the eastern hemisphere were primitively

BEB (ALES (00 Eee PR EB CM ae oat Ce Voy ISIS sae sve. w) si < va o lols “csi dl avoe fhe lbes wialel'e cote 80

TUB SIRES ois wats BSc b esis ET SR Ae eee ore ca 80

CTISTACEA Ha sss a eee arena ciel eave ol ncaa Sau aeparapoes ei 80

AMOUR 5 5 ois oS agile Sete egg a ete A le ei ee 80

BATES > oo 5b sain Sieve Sled GS OTE Oe en Me aE ae ea 83

OPES s cob at UPR SOR Seo eee eee ann 83°

(GOMER TOPS. ere po B Cues PCO ean epee 84

FoTON AUB ILE TO ILCSIBMER IEP gS cron exe Siena neky cat hays cc 2 chad algvapiacsasd dev ately quale 95

NHN 5 5,4 WG a be Bee OU OS GP ENOL Sa en et ae rr a irra 99

Scag SRN Tell Te PP eee ee eh se le dase x cpociaivv'c o 6 u'o-bace® ceetsnecuwed 103

TES ONO BYINY 5555.9 BIS aS Oh cr Ot SI ete as es i 106

10 ANNALS NEW YORK ACADEMY OF SCIENCES

INTRODUCTION

South America has not lacked the labor of scientific explorers; none

the less one may safely state that fully one half its surface is still prac-

tically unknown. This is due to the fact that its investigators have been

relatively few, but even had there been many scores of them they could

not adequately have explored such an expanse of land beset with count-

less natural obstacles.

When attention is drawn to the fact that Brazil, which is one of the

best known South American republics, is larger than the United States

itself, it becomes evident that the few dozens of explorers within recent

times could not have examined more than the regions accessible from

the coast, from the few railroads and from the !arger rivers.

Pioneer work in the interior of tropical America must often depend on

meager and often incorrect information. This is reflected even in the

maps of South America, for in every one some of the material has been

taken from untrustworthy local sources.

It was on account of this lack of decisive knowledge concerning cer-

tain parts of South America, even in the best books of reference, that the

director of the Carnegie Museum dispatched the writer, in 1907, in

charge of the Carnegie Museum Expedition to Brazil. One of the

primary objects of this journey was to study the distribution of the

fishes; but kindred problems were not to be neglected, and data were

collected on every hand and over a far greater territory, partially known

or quite unexplored, than had been originally suggested. This was found

possible because the writer was fortunate in maintaining excellent health

throughout his journeys. (See Plate II for routes followed.)

For this excellent opportunity to explore many regions of South Amer-

ica, [ am deeply indebted to the founder and the trustees and to Dr.

W. J. Holland, the director of the Carnegie Museum.

Acknowledgment should be made to Dr. O. A. Derby, the director of

the Brazilian Geological Survey, for many favors and much useful in-

formation in furtherance of my work; to Prof. J. C. Branner and to

Mr. R. Crandall, both of whom assisted me notably on my first trip to

Bahia. Mr. Crandall examined my notes and map of the Cretaceous

*of northern Brazil. This enabled me to add valuable corrections. He

has also kindly submitted me some notes on the trend lines of northeast-

ern Brazil. Dr. Schuchert has put me into his debt for many invaluable

suggestions, and so too has Dr. David White for corrections and sug-

gestions in the discussion of the Gondwana flora.

My thanks are alse due to the directors and staffs of the museums at

'——<-

ANNALS N. Y. AcApD. Scr.

VOLUME XNIJI, PuatTe II

dnt pagent

Bhonsafnaslatin

fopese

Tonal

Celdara

Fon te! Moae

Vova vier

MAP OF PART OF SOUTH AMERICA

Showing journey made by J. D. Haseman under the auspices of the Carnegie Museum,

1907 to 1910, in Brazil, Uruguay, Argentina, Paraguay and Bolivia

HASEMAN. GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 11

Rio de Janeiro, Sao Paulo, Para, Montevideo and Buenos Aires, and

among my other South American friends, to Ricardo Krone of Iguape,

John Gordon, Alipio Miranda Ribeiro and Carlos Moreira of Rio de

Janeiro, Dr. Jappa Assu of Bahia, Rudolfo von Ihering of Sao Paulo,

Dr. Frank Davis ani Feliciano Simon of Corumba and Dr. Snetlager of

Para.

Among my preceptors, I record gratefully my indebtedness to Dr.

Higenmann, with whom I carried on my special studies in the University

of Indiana; to Dr. A. E. Ortmann while I was in Pittsburgh, and to

Drs. Dean, Grabau. Gregory and Hussakof while at Columbia, where a

scholarship and the income of the Dyckman Fund for 1910 were gener-

ously granted me.*

The living and extinct fauna and flora of South America possess in

certain cases, at least, a close genetic relationship with that of the south-

ern portion of the 2astern hemisphere. This remarkable fact has led to

a prevailing view that South America was cnce connected with some

point of the eastern hemisphere. This view assumes that the closely

related fauna and flora are descended from common ancestors which

existed in the old land mass or continent to which the name Gondwana

has been applied.

In this thesis, it will be my effort to show that at least certain, if not

all, elements in the fauna and flora of South America have evolved from

forms which have from time to time been introduced from the northern

hemisphere. In this view, I have brought together not only materials

from references and from the laboratory but also data obtained during

two and one half years of active field work. The amount of this ma-

terial altogether will be sufficient, I think, to demonstrate that South

America was never connected with the eastern hemisphere by a hypo-

thetical southern and sunken mass of land.

In the preparation of my paper, I have been obliged to omit numerous

data and assorted faunal lists whose bearing has been more or less direct

upon the present theme—in the latter cases since I am convinced that

the lists do not explain distribution unless accompanied by detailed ob-

servations upon the geology and the environmental conditions of the

country considered. Hence I have been led to divide my thesis into two

parts. In the first of these, we picture the past and present environ-

ments in which the fossil and existing animals lived. In the second, we

deal with the changes through which these animals and their ancestral

stocks have undergone after arriving in these environments.

1The geographical names are spelled as they are in their respective countries. I have

also used the old way of spelling such words as “Silurian” instead of “Siluric.”” I have

also omitted marks of accentuation and other similar marks.

_—

ANNALS NEW YORK ACADEMY OF SCIENCES |

It is in the first part of this thesis that my effort will be to demon-

strate on geological grounds that South America has not been connected

with the eastern hemisphere. In this connection, I have also been able

to map for the first {ime the outline of the Plano Alto (the highlands of

South America) which was deposited by the wind and rivers in a dry

land and fresh-water basin which I am calling the Permian Inland Basin.

This highland, I propose to show, was of the utmost continental impor-

tance, from its dip, its lack of Mesozoic and Tertiary marine deposits and

the direction of the trend lines, taken in connection with the Tertiary rise

of the Andes; in fact, upon this I shall base my doctrine that the Amazon

is a reversed river whose headwaters originally flowed into what I have

designated the East Andean Sea.

This outlining of the Plano Alto is of prime importance, not only be-

cause it has given me the key to the correct explanation of the distribu-

tion of the aquatic life, but also because it shows that South America has

not been cut into islands by east and west invasions of the sea as has

been proposed by some of the exponents of the Archhelenis theory of

von [hering.

In other topographical matters which are of importance from the

zoogeographical viewpoint, I will also show that the Paraguay River is

not connected with the Guapore, as has been so often erroneously stated,

and that Rio Sao Francisco is connected with Rio Tocantins. I will

note additional cases of stream piracy and will show that these taken in

connection with waterfalls, swamps and certain environmental conditions

will aid us in interpreting with a certain degree of accuracy questions in

the distribution of the South American fishes. :

In the last part of this thesis, an attempt will be made to demonstrate

that the fauna of South America has been evolved from the forms which

originally lived in North America. In arriving at this conclusion, I

have not entirely limited my studies to the fishes, but I have considered

carefully the voluminous data derived from other groups of living and

extinct animals and plants which have been used to establish connections

between South America and the eastern hemisphere. I have considered

all of these data, because the facts derived from the distribution of any

one group of plants or animals are not sufficient alone to warrant an in-

terpretation which involves profound modifications of the earth’s surface

as maintained by many authors.

In the matter of the distribution of fishes, as bearing upon the greater

problem, my effort will be to show

1. That the present distribution of fishes gives no clue to the point of

origin of the families, but it does for some of the genera and many species.

2. The point of family origin can only be determined after the living

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 15

and extinct forms have been carefully compared and a sharp distinction

is made between paleotelic (old or phylogenetic) and cenotelic (recent,

adaptive or physiological) characters. It is necessary to draw this dis-

tinction, because only paleotelic characters have been widely distributed.

3. That a fresh-water connection or its absence will not alone explain

the present distribution of the fishes. Hence, the most important factor

of living fish distribution is not land and water connections, 7. ¢., bar-

riers, isolation, intermingling, etc., but it is the organic complex of the

ancestral stock and the effects of different environments on this stock.?

4. That much of the similarity and some of the identity of certain

species of the fishes of Rio Paraguay and Rio Amazonas are due to simi-

lar and identical evolution of the highland ancestral stock after arriving

in similar environments, e. g., as produced by the erosion of the high-

lands.

5. That the existing highland genera, small in size, are the more gen-

eralized types from which the bulk of South American fishes has evolved.

6. That the South American fishes have evolved from primitive forms

which originally lived in North America.

The above statements do not in the main agree with the views ex-

pressed by previous zoogeographers. This difference of opinion is largely

due to the fact that these investigators work from the static viewpoint of

animal geography and have therefore only considered in some cases the

“disconnected graveyard material,” i. e., a few isolated spots where the

fauna died out; and in other cases the “hot-bed material,” 7. e., the end

result of the greatest cenogenic evolution. For this reason, their static

faunal lists do not correctly determine the point of origin of families

and orders. As a result of their conception of animal geography, some

of these writers have maintained invasions of the sea and land-bridges

for which there is no evidence. They have also brought to the support

of their views some unnatural environments and unwarranted views of

the geology and the topography of South America.

Another source of error in former interprctations is, I believe, the

ignoring of the possibility of similar evolution of the identical ancestral

stock in remote but similar environments. The necessity for the recog-

nition of such evolution is due to the fact that in the same river basin

there often exist (two or three) distinct fau.a! regions, one of which

may show close affiuity with another distinctly separated basin, while

another river system, although connected (with the latter), may yet re-

tain quite distinct faunas.

2The idea that isolation alone produces new species implies the principle of selection,

which is still of doubtful value. If it is not selection, then it must be in some way the

direct or indiréct (also debatable) influence of the environment on the germplasm or

else it is orthogenesis.

14 ANNALS NEW YORK ACADEMY OF SCIENCES

My studies on the distribution of South Aierican animals have also

led me to place more emphasis on negative evidence than is usually

granted by most writers. This difference of view is primarily due to the

fact that a specialist in any one group of animals places too much weight

on his positive evidence. Such emphasis at iirst sight appears to be

absolutely correct, but on closer analysis it is usually contradicted by

positive evidence from other groups of animals. For example, one au-

thor working on crustacea and mollusca finds that the alleged connection

between South America and Africa had already disappeared in the early

Cretaceous. If, however, we consider other groups, we find evidence

which does not confirm this view. Thus, the affinity existing between

the South American and African characinid and cichlid fishes is as close

as that of the mollusca, yet there is no evidence that either of these

families of fishes existed during the early Cretaceous.

In my conclusions, therefore, I have been led to balance the positive

and negative evidence in the cases of many different groups. This bal-

ancing has been attempted not only by considering long lists of species,

but by taking into due account the influence of various environments on

the ancestral stocks (whose points of origin are usually unknown).

In fact, we should not, in such considerations, lose sight of the fact

that our knowledge of the existing species of any group of South Ameri-

can and African animals is still very imperfect. Many species are still

to be caught, many are exact synonyms and many are without doubt

local somatic changes which are not always inherited. Therefore any

positive evidence derived from such lists is, in my opinion, entirely in-

adequate to warrant the reconstruction of the earth’s surface, unless

supported by strong geological evidence. ‘This is all the more true when

there are other means of distribution which do not involve great topo-

eraphical changes.

In point of fact, the inadequacy of the Seine data is at once re-

flected by the number of alleged land-bridges and seas required during

various past geological epochs. Each writer has constructed his new set

of barriers, seas and land-bridges in his effort to explain the distribution

of the fauna or flora in which he is recognized as an authority.

Notwithstanding the diversity of views concerning the time of exist-

ence and the location of the alleged land-bridges to the South American

continent, we may roughly consider them under the following two

groups:

1. The Gondwana Land of Suess and others—a late paleozoie conti--

nent traversing the greater part of the southern hemisphere and connect--

ing India, Australia, South Africa and South America. The last re-

mains of this old land-mass connecting Africa and South America have

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 15

been designated Archhelenis by von Ihering. Antarctica, another por-

tion of Gondwana, is the name generally applied to the south polar conti-

nent which is believed by some to have been connected with South

America and Australia and perhaps with Africa.

2. The persistence of the continental shelves and the great ocean

basins. This view precludes the existence of furmer connections between

South America and the eastern hemisphere, but admits the North Ameri-

ean connections with Eurasia.

These two views involve not only the distribution of plants and animals

but the geology of the entire earth. They are not new, dating back in fact

nearly to the time (1857) when Sclater first placed geographical distribu-

tion in tangible form. In his scheme of distribution, the world was con-

sidered in six faunal regions, a scheme in which South America formed a

major part of the neotropical realm.

_In 1876, Wallace published two comprehensive volumes on the geo-

graphical distribution of animals. In these two most excellent volumes,

the genus was used more than the species as a means of comparing faunal

regions, and it appears that such a comparison is more luminous than one

based on the species, for it is found that the generic characters are usually

more nearly paleotelic than the specific ones; moreover, the list of species

is the less accurate, since a far greater number of species than genera are

still undescribed. The extensive data given in these two volumes indicate

that the bulk of the ancestral land animals originated in the northern

hemisphere. It is worthy of note that the views expressed by Wallace and

some of the other earlier writers were far more conservative than those of

more recent date.

It was during the interval from 1876 to 1890 that zodlogical, paleonto-

logical and geological data accumulated rapidly and yielded, especially,

the excellent summation of the geology of the face of the earth by Suess

and his views of the Gondwana Land (earlier suggested on purely paleon-

tological grounds by Neumayer), and his considerations appear to have

paved the way for von Ihering’s Archhelenis theory which has more or

less dominated most of the later studies on the zodgeography of South

America.

Von Ihering has made, from time to time, slight changes in his theory,

in order to meet the demands of more recent investigations. In 1907, he

reconstructed the surface of the earth according to the views which he

obtained from a detailed study of the mollusca. At the time, he main-

tained that, previous to and during a part of the Tertiary epoch, Brazil

(Archibrazil or Archamazonia) was connected by Archhelenis with Africa

and by Archiplata with Archinotis (Antarctic continent), which was also

16 ANNALS NEW YORK ACADEMY OF SCIENCES

continuous with Australia. The sea traversed northern Brazil and sepa-

rated the Guiana highlands plus the West Indies (Archiguiana) from the

rest of South America, but it was connected with both Asia and Europe.

Africa was not connected with Asia at that epoch. In 1911, he had made

many changes in his views. P

On the other hand, after a long detailed study of both the mollusca and

crustacea, Ortmann has also maintained that an Archhelenis existed, but

he differs with von Ihering both in regard to its location and the time of

its disappearance. He believes that Archhelenis had already disappeared

before the beginning of the Tertiary (perhaps the early Cretaceous) and

that it connected Guiana and Africa.

Eigenmann (1909) has tested the Archhelenis theory with the distribu-

tion of the South American fishes and has found no objections to it. In

fact, he states that the theory is quite useful in explaining the distribution

of certain families of fishes, especially the Characinide and the Cichlide.

D. White (1907) in an excellent paper on the Gangamopteris flora

(Gondwana flora) of Brazil does not, however, favor the Archhelenis

land-bridge; he believes that an ancient connection (Permian) very

probably existed between South America, the Antarctic continent and

Australia or Africa.

In Part V of Volume III of the Princeton Patagonia reports, Pilsbry

has summed up the distribution of non-marine mollusca of South Amer-

ica. His figure, page 632, indicates a former connection between the

region of Pernambuco, Brazil and South Africa. He is inclined to believe

that this connection disappeared by the end of the Cretaceous. He also

admits the probability of a connection by way of the Antarctic islands

with Australia, but he does not believe in a former isolation of the Guiana

highlands from those of Brazil.

Schuchert (1911) also believes that the distribution of the brachiopoda

shows clearly not only the former existence of an equatorial Gondwana

across the Atlantic, but as well that its vanished Atlantic bridge still con-

trols the distribution of living forms. He is of the opinion that Gond-

wana probably existed until middle Eocene times.

In the Age of Mammals (1910), Osborn re-states the widely accepted

belief in an Antarctic connection between South America and the Aus-

tralian realm, but he rejects a Tertiary Archhelenis. He thinks that this

connection is necessary in order to explain the great similarity which

exists between some of the fossil marsupials of Patagonia and the mar-

supials of the Australian realm. He also states that Matthew has rejected

both the Archhelenis and the Antarctica connections and now maintains a

northern origin of the southern fauna.

ANNALS N. Y. ACAD. SCI. VOLUME XXII, PLATE TEI

a EE ER A EC LET

ARCHEAN AND OTHER PRE-CAMBRIAN AREAS OF SOUTH AMERICA

The solid black represents the Archean; thee horizontally lined area represents the

basal highland formation, which is generally considered to be pre-Cambrian and which

is covered by alluvial deposits along the Amazon (dotted area).

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 17

The view held by Dr. Matthew and the writer (who have independently

and from different standpoints arrived at several identical and funda-

mental conclusions concerning the distribution of South American ani-

mals) in a general way was put forth in 1886 by Haacke as the North

Polar theory of the origin of land animals.*

Part J. GroLtocy anp TopoGraPHy oF SourH AMERICA #

DISTRIBUTION OF GEOLOGICAL HORIZONS

It has been extremely difficult to map even the larger divisions of the

geological ages found in South America, since the exact age and extent of

many of the known formations have never been satisfactorily determined.

Archean

Archean rocks, as shown in a general way in Plate III, extend in a nar-

row belt, more or less broken, from Tierra del Fuego to the Isthmus of

Panama. A great depression east of the Bay of Arica hes between the

coastal Archean rocks of Chili and the inland Archean rocks of the north-

ern Andean region. Another great belt of Archean rocks extends from

Uruguay into the Serra do Mar and its various northern spurs of eastern

Brazil. Archean rocks have also been encountered in isolated places of

Patagonia, and a belt more or less broken extends from near Bahia Blanca

through the Cordova Mountains into southern Bolivia. Finally, rocks of

similar age have been encountered in northern Colombia, parts of Vene-

zuela and Guiana.

The exact age of the crystalline schists, gneisses, granites, etc., which

‘underlie the Plano Alto has never been satisfactorily determined, but they

are usually considered pre-Cambrian or Archean on account of the entire

absence of fossils. The lined portion of Plate III indicates the extent of

these rocks.

Lisboa, one of the most competent Brazilian geologists, has reported

crystalline rocks of pre-Cambrian (?) from near Miranda, Matto Grosso

and in the basins of Rio Apa and Rio Aquidauana. The writer observed

the basal highland rocks on the Bolivian side of Rio Guapore below the

mouth of Rio Verde and on the Brazilian side of the Guapore at the

2 The bibliography gives a list of the publications which have been extremely useful to.

me, and I take the opportunity here to acknowledge my indebtedness to the authors. Few

references have been given in the pages of this thesis, because it has been deemed ad-

visable to omit them for the sake of clearness and brevity. Therefore some common

information has been freely used.

+In the preparation of this part of the thesis, the writings of Derby, Branner, Suess,

Eschwege, Hartt, Hatcher, Steinmann, Phillipi, Stelzner. Hauthal. Katzer, Crandall and

numerous other authors given in the bibliography have been indispensable.

18 ANNALS NEW YORK ACADEMY OF SCIENCES

waterfalls of Forto de Principe da Beira. They were also seen along Rio

Mamoré above Guaja Mirim, where the Serra de Pacas Novas approaches

the river. Evans has reported gneisses, etc., from the Rio Beni and the

Madeira falls. WKatzer has maintained that this same basal highland for-

mation extends under the deep alluvial deposits of the lower Amazon

Valley. Asa result of the writer’s own observations, he is convinced that

this is in the main the correct view. Hence this basal highland formation

is very large, perhaps larger than indicated on Plate III.*° The extent

and form of these ancient rocks have given South America its present

shape from the very beginning.

Silurian ©

Silurian fossils have been reported from Rios Curnua, Maecurt and

Trombetas, which are affluents from the north side of the lower Amazon

River; from Bom Jesus da Lapa, of Bahia; from some of the promon-

tories of Venezuela; from the eastern chains of the mountains near the

headwaters of Rio Bermejo, Argentina; west of San Juan along the

eastern base of the Andes of northwestern Argentina; from Sierras de

Famatina and La Rioja; from the mountains on either side of Sierra de

Aconquija; from Sierra Aguilar, and from Cuzco, in southern Peru, ex-

tending past Hlampu and Illimani through western Bolivia toward the

Argentine chains of mountains. (?) Silurian (Arthrophycus harlant)

was reported from Sierra de Ja Tandil.

The locations of these formations are shown on Plate IV, except ‘that

no distinction has been made for Ordovician because of the lack of data.

Devonian

Devonian fossils have been reported from Alameirim to Rio Uatuma at

Erere, Rio Maecurt south of Larangal on the south side of Rio Amazonas ;

from Lagoinha near Cuyaba; from a belt extending from the State of

Sao Paulo into the State of Parana at Ponto Grosso, at Jaguarahyva and

in the Ivahy basin ; in southern Peru and about Lake Titicaca, and in the

Andes of Chili. Doubtfully from the northern part of Sierra Tandil.

The locations of these formations are shown on Plate V.

Carboniferous

Carboniferous fossils have been reported from Rios Trombetas, Curua,

Maecurt, Uatuma to Janary near Prainha and at Alemquer on the north

5 It must be granted that the regions supposed to yield exposures of Archean rocks may

with more careful study be materially diminished, as has been the case in North America

and Europe. If these rocks mapped as Archean are all older than the Carboniferous

formations, then the ensuing views are not in the least affected.

6 The Cambrian has not been mapped because of the lack of data. Deposits of this age

have only been reported from a few places, as northern Argentina by Keyser, etc.

ANNALS N. Y. ACAD. SCI. VOLUME XNII, PLatTe 1V

of, <\

ENOWN MARINE DEPOSITS OF SILURIAN AGE IN SOUTH AMERICA

i¥

ANNALS N. Y. AcapD. Sct.

2—_NY

VOLUME XXII, PLaTse V

: ism

of ~

en LP

SON

KNOWN MARINE DEPOSITS OF DEVONIAN AGE IN SOUTH AMERICA

ORT at

ANNALS N. Y. Acab. Scr. VOLUME XXII, PLatE VI

ie WSS

pu? )

KNOWN MARINE DEPOSITS OF CARBONIFEROUS AGE IN SOUTH AMERICA

ce AE ey

aaa tea

ANNALS N. Y. ACAD. SCI. VOLUME XXII, Pratn VII

A) 2g

y (

< OP

NY us

KNOWN MARINE DEPOSITS OF PERMIAN AGE IN SOUTH AMERICA

ANNALS N. Y. ACAD. SC1. VOLUME XXII, PLaTE VIII

KNOWN MARINE DEPOSITS OF TRIASSIC AND JURASSIC AGE IN SOUTH AMERICA

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 19

‘side of Rio Amazonas and along Rio Tapajos from Itaituba to Aveiro and

west to Maueassu at Fructal and Rio Pedra do Barco on the south side of

Rio Amazonas; from the Cordillera Oriental of Peru; from Lake Titicaca

near Yampopata and extending south towards Cochabamba at Arque,

Bolivia; Santa Cruz, Bolivia, and in the Chapoa Valley at la Ligua,

Chil. The locations of these formations are shown on Plate VI.

Permian

Permian fossils have been reported from the State of Sao Paulo, Brazil,

and this same belt extends south through Parana and Rio Grande do Sul

and ends in Uruguay. Permian fossils are also known from a few isolated

places in the Andes. The location of these formations is shown on

Plate VII.

Triassic and Jurassic

The Triassic and Jurassic periods are unknown in Brazil. ‘Triassic

and Jurassic fossils have been reported from northern Colombia and

Venezuela; from Puerto Puruay and Rio Maranhao of Rio Amazonas,

and are widely spread in the Andes of Peru and Chili at Passo de los

Patos, Coquimbo, Copiapo and elsewhere, where they are mixed with fresh-

water deposits. The locations of these formations are shown on Plate

VET.

Cretaceous

Cretaceous fossils have been encountered in a narrow belt along the

Brazilian coast from Ilhéos, south of Bahia, to Pirabas, near the mouth

of Rio Amazonas. This coastal belt is never more than about 100 feet

above the sea level, and its fossils also show that it must not be confused

with the inland Cretaceous of northern Brazil which extends north from

the State of Bahia across Rio Sao Francisco at Jatoba and widens out in

the states of Piauhy and Ceara. This belt is often 1000 or more feet

above the sea level. If the writer is correct in considering a formation

(in which he found no fossils, but near where at Curimata fossil fishes,

(?) Diplomystus, are said to exist) near Lagoa da Paranagua in the

State of Piauhy as Cretaceous, then this belt will with all probability

later be found to extend as far south as Serra da Tabatinga, but it does

not extend into the Jalapao region of northern Goyaz.

The extreme northwestern extension of this Cretaceous is unknown,

and consequently it offers alluring opportunities for future study,’ be-

77This belt may extend northwest into the State of Para and as far west as the lower

‘Tocantins basin. Mr. Roderic Crandall of the Brazilian Geological Survey has done the

most field work on this inland Cretaceous belt, and his views agree in the main with

those of the writer.

20 ANNALS NEW YORK ACADEMY OF SCIENCES

cause it will extend the limits of the Cretaceous deposits over a vast

portion of northeastern Brazil.

Cretaceous deposits have been reported from Bahia, Espirito Santo,

Aracaju, Alagoas, Maria Farinha, Jatoba, Riacho Doce, Serra de Ara-

ripe and Pirabas in Brazil; from the base of the mountains of Guiana;

from Bogota and the region of Lake Maracaibo; from Sierra de Merida

of Colombia and Venezuela; from Cordillera Nevada and from both

sides of the Andes in the region of Alto Rio Maranhao; from Rio Acre

(Mosasaurus) ; from Caracoles, Bolivia; from near Lima, Peru; Cochi-

yacu west of Rio Huallaga and north to Celendin; from near Guayaquil ;

from Tingo and south along both sides of the Andes toward Chili at

Tomé, etc.; from Laguna Argentina to Tierra del Fuego; from Rio de

los Patos west of San Juan, Argentina; from Colchagua, Coquimbo and

Copiapo in Chili; from Sierra de Zenta east of San Juan, and perhaps

from Gran Chaco toward the mountains about the headwaters of Rio

Bermejo and Pilcoinayo, but I am inclined to believe that these de-

posits are fresh water, as the most common form, Melania, is not typi-

cally marine. At any rate, this region needs some more careful study.

The locations of these formations are shown on Plate LX.

Tertiary

Professor Branner is inclined to consider some of the marine forma-

tions of northeastern Brazil Eocene. This view has been recently cor-

roborated by President Jordan’s studies on the fossil fishes from Riacho

Doce, but neither of these authors has entirely excluded the possibility

of these formations being upper Cretaceous.

As far as the writer has been able to ascertain, from a first-hand

knowledge of the region in question as well as from that of Crandall

and also from a consideration of President Jordan’s paper on the fossil

fishes of the Serra de Araripe, it does not appear that any decisive evi-

dence exists which establishes any marine Tertiary in northeastern

Brazil. Fossil diplomystid fishes, the subject of President Jordan’s paper,

are known not only from the Cretaceous of Brazil, but also from the

Cretaceous of other Continents. The fact that most of the diplomystids

are found later than the Cretaceous epoch is no evidence that those of

Serra de Araripe are Eocene. Furthermore, the peculiarities of the

diplomystids of the Serra de Araripe will, with all probability, be found

in various other localities of this region, when more exploration is com-

pleted.

For several reasons, therefore, I have mapped the outline of the Cre-

taceous belt on the map of the Tertiary epoch with a mark of imterro-

ANNALS N. Y. ACAD. SCI. VOLUME XXII, PLATE IX

KNOWN MARINE DEPOSITS OF CRETACEOUS AGE IN SOUTH AMERICA

ANNALS N. Y. ACAD. SCI. VOLUME XXII, Pratn X

U \

4 \ (

‘ \ .

' \ ~

1 \ On

, ‘ .

‘ i, ~s

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( ee ‘

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‘ . 1

a oN y

sien i ’

és a

eer

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‘

KNOWN MARIND DEPOSITS OF TERTIARY AGE IN SOUTH AMERICA

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 21

gation, for later work may establish Tertiary ceposits in at least part of

this region, especially if we accept Professor Branner’s view of its

stratigraphy.®

Marine Tertiary fossils have been reported from Entre Rios, the Pam-

pas and part of the Gran Chaco of Argentina; from the eastern base of

the Andes, extending into the plains of Patagenia and as far south as

Tierra del Fuego; from narrow belts along the coast of Peru and Chili;

from the lower Orinoco valley; from Pebas, Peru, down Rio Solomoes as

far as Sao Paulo de Olivenca, and from Canama on Rio Javary and prob-

ably from the region of Rio Acre, Brazil, and Santa Maria, Catamarca,

west of Jejuhy in northern Argentina. The locations of these formations

are shown on Plate X.°

TREND LINES

The trend lines of South America are about parallel to the coast, ex-

cepting in the region of Pernambuco and Ceara, Brazil. In this region,

they appear to be fan-shaped. In regard to this most interesting region,

I can do no better than quote a letter from Mr. R. Crandall, of the

Brazilian Geological Survey, who has explored this region during the

past four years:

“The general trend lines through all of the State of Bahia, as you will re-.

member, are northwest-southeast and north into northeast-southwest. These

lines get lost as we get farther north, and the trend of the coast itself changes

north of Pernambuco. The change in the trend of the coast is accompanied by

a Similar change in the direction of the Serra da Borborema. The Serra da

Borborema is more properly an eroded mass than a structural line, though it

conforms quite closely to the general trend of this region.

“I consider the lines in the Ceara, Rio Grande and Parahyba true structural

lines, as they are Jong lines of intruded granites and allied or similar rocks

which indicate intrusions on or along lines of previous weakness. Part of these

lines are indicated by the Ceara series of rocks which I have correlated with

the Jacobina series of Bahia and with the Minas or iron-bearing series of Minas

Geraes.

“T have never properly understood the forces that formed this fan-shape in

northeastern Brazil. Just at what age it came is hard to say, as the age of the

latest folded rock, the so-called Ceara series, is about Cambrian (for all we

know, even pre-Cambrian). I believe that Derby considers the granites of

southeastern Brazil to be post-Devonian, and I believe that these northern

SIt does not make any great difference, as far as the present conclusions are con-

cerned, whether marine Tertiary does or does not exist in the above region. Professor

Derby (1907) also expresses some doubt about the existence of marine Tertiary in the

said portion of Brazil.

®Tt is beyond the scope of this thesis to attempt to discuss in detail any of these for-

mations, excepting that of Alto Rio Amazonas. The necessity of this preliminary sketch

will become evident after the reader has considered the ensuing topics. What has been

land and sea is important from the standpoint of animal geography.

ae ANNALS NEW YORK ACADEMY OF SCIENCES

granites are in the larger part of an age younger than the Ceara series and.

somewhere along in the Devonian is entirely probable. You see between the

Cambrian and the Cretaceous in the northern region we have no record.

“Tt is entirely possible that the northern fan is due to local folding, but it is.

pretty large for that, that is to say, spread over a very large area.”

I believe that these facts indicate that no Paleozoic wedge or land-

bridge could have existed between this portion of Brazil and Africa,

because the trend lines fade away toward the sea, and there is no eyi-

dence of the continuance of the lines across the Atlantic into Africa.

The folds are crushed and irregular and do not end abruptly along the

coast. Besides, if the wedge had existed, this type of fanlike folding

would, I believe, have been almost if not quite impossible. This fan--

hike structure may possibly have been produced by some unknown force:

pushing from the interior of the earth at an angle to the vertical and.

more or less parallel to the coast, but more strongly in the region of the

mouths of Rio Amazonas and Rio de la Plata, if the Brazilian coast was.

not connected with Africa. This would have pushed the southern and.

northern ends of South America to the west and have made the “Per-

nambucan fan” on the east and a somewhat similar structure in the region

of Lake Titicaca east of the Bay of Arica.

In southern Brazil and more especially in val of the Plano Alto, ero-

sion and extensive contine::tal deposits have so disfigured the unexplored.

forested surfaces that littic is known concerning the trend lines, but:

superficially this region looks like an abruptly chopped-off coast. This

abruptness, I believe, is due to erosion of late and post-Paleozoic land

deposits and not to post-Paleozoic faulting. The northwest strike from

Cuyaba past the Madeira Falls, noted by Evans may be due to ancient

erosion, but I hardly think so, because the Plano Alto dips toward the

southwest. The old drainage was into the Hast Andean Sea and hence

the rivers at that time cut the Plano Alto im a western-southwestern

direction. When the Amazon became reversed, Rio Madeira cut these

old planes of erosion almost at right angles, producing thereby a very

complicated topography.

By means of the trend of the Sierras de Tandil and de la Ventana, we:

can separate both of them from the Andean complex, in spite of the fact

that Suess and Stelzner have been inclined to consider the Sierra de la.

Ventana as belonging to the Andean system.

In fact, the writer considers that the Sierras de la Ventana and Tandil

with all of their side chains are the southern extension of the Cordova

Mountains by the way of Sierra de San Luis, where there is a break in

the system, just as the Serra do Mar of Brazil breaks up into several

es ~

— =

ANNALS N. Y. Acad. Sci. : VOLUME XXII, PLatTE XI

OUTLINE MAP OF SOUTH AMERICA

Showing the trend lines about the Plano Alto and the hypothetical outlets of the Hast

Andean Sea. The “Pernambucan Fan” has been put in from notes and maps contributed

by R. Crandall.

3—NY

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 23

chains in east central Brazil. I also believe that the above sierras origi-

nally formed part of the southern and western boundary of the great

Permian Inland Basin which will be considered in the following pages.

I have relegated both the Sierras de Tandil and de la Ventana (as

Hauthal did several years ago) to the Brazilian system for the following

reasons :

1. The Sierras de Tandil, de la Vantana, de San Luis and de Cordova,

like the Serra do Mar of Brazil, are characterized by an almost entire

absence of marine fossils and have no marine Tertiary and Mesozoic,

which is characteristic of the Andean complex.

2. In general succession; the Archean rocks of these sierras resemble

the Serra do Mar rather than the Andean system.

3. These sierras are well separated from the Andean complex by

many elevations and depressions, some of which afford evidence of Meso-

zoic and Tertiary invasions of the sea.

4. From the absence of fossils, it may be judged that these sierras

have remained almost stationary; and as a result, there is a striking

correspondence between their altitudes and these of Brazil and Guiana.

It is possible from similar reasons that at least part of the Cordilleras

Oriental of the northern Andes will be shown to belong to the old Bra-

zilian system, but entirely too little is known about this region to war-

rant a consideration at this time.

The trend lines of the Pacific side are also parallel to the coast. In

the region of La Paz, Bolivia, there is an indentation of the coast back

of which the chains of mountains are bent out of line and piled up to

several thousand feet of altitude. This region is, in a way, the counter-

part of the fan-shaped region of Pernambuco along the Atlantic Coast.

In other words, the trend lines of the two coasts exhibit a remarkable

similarity, as is indicated in a general way on Plate XI. The only strik-

ing difference is thay in the region of La Paz, Bolivia, the coast is bent

in, while it is bulged out in the region of Pernambuco. ‘Taylor has

attempted to explain part of this by a sliding of the Brazilian mass

against the Andes. This sliding was assumed to be due to some force

applied parallel to the Brazilian coast but having a greater intensity in

the regions of the mouths of the Amazon and !a Plata.

The lines of weakness and strength in a general way extend north and

south (as Schucheri has shown for North America). This is shown by

the maps of the marine deposits. The invasions of the sea as a rule

appear to have been from the south toward the north. The Permian of

southern Brazil or the Devonian extending from the Amazon Valley

past Cuyaba into Parana, and perhaps as far south as Sierra de Tandil,

24 ANNALS NEW YORK ACADEMY OF SCIENCES

if the (?) Devonian of Siemiradzki exists there, are good examples of a

southern-northern invasion of the sea. In all of this region, an east to

west invasion of the sea appears to have been impossible, excepting in

narrow belts along the coasts, on account of the intervening Archean

mountains, like Serra do Mar, which show no traces of marine deposits.

This is of the utmost importance from the standpoint of animal dis-

tribution. Some authors have attempted to show that the Patagonian

region, and others that the Guiana region, was for a long time cut off

from Brazil by arms of the sea. In order to isolate either of the above

regions from Brazil, it would require an extensive east to west invasion

of the sea for which we have no evidence. On the other hand, the maps

showing the location of marine deposits offer strong evidence against

such a view. Hither the older rocks of Chil or of the Cordova Moun-

tains could afford connections between southern South America and the

Brazilian region. Uence the observed difference in the fauna of Pata-

gonia must be due in a great part to environmental conditions.*°

. BRAZILIAN COAST

In some respects, the Brazilian coast appears to be the counterpart of

the contour of West Africa. Its abruptness is thought by many to be

due to the submergence of a “wedge” which originally connected Brazil

and Africa.

Soundings have shown that deep sea exists within a comparatively

few miles of the Brazilian coast; but thus far soundings have not pro-

duced the slightest evidence for a submerged “wedge” or land-mass

which is believed by many to have originally connected Brazil and

Africa. In fact, there is strong evidence derived from soundings against

the submergence of such an extensive land-mass into the abysmal depths.

This wedge must have been deeply eroded, forming thereby deep, wide

and abrupt valleys. When this surface (sucl as Brazil at the present

time) dropped beneath the ocean, few soundings would be needed to show

that the bottom of the ocean under such conditions would not be uni-

form. Inasmuch as soundings have revealed no evidence in favor of such

a rough sea bottom, | take this as strong evidence against such a view.

It is true that Murray has found a mid-Atlantic ridge, but the trend

of this elevation is parallel to the distant coasts, 7. e., more or less north

and south and not east and west.

10 Also the imperfection of the fossil records, exploration, etc., in South America can-

not be ignored. I saw part of a Torodon from near Uruguayana, Brazil. Even if an arm

of the sea separated Patagonia from the rest of South America, it would have been

entirely too narrow to have been an effective barrier.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 25

Soundings as well as fossils indicate that the Brazilian coast at one

time extended slightly farther east than at present. There appears

to be little or no doubt that the Abrolhos and Barbados islands were

originally connected with South America (? late Mesozoic). To the east

of these islands, no shallow sea or numerous small islands exist, as they

do in regions like the West Indies, East Indies and Alaska, where we

know that elevations and subsidences have taken. place.

The abruptness of the present Brazilian coast is not necessarily evi-

dence in favor of a post-Paleozoic faulted coast.. In fact, my own ob-

servations have convinced me to the contrary. The entire absence of

fossils along the coast of southeastern Brazil indicates a great stability

of this region. Inasmuch as in all this region nearly all of the rivers

flow west and southwest, away from the ocean, it appears that there is a

gentle dip towards the southwest. Of course much of the surface of

this region is covered by late and post-Paleozoic continental deposits

which have been deeply eroded. The erosion of these deposits as well

as the older rocks would produce an abrupt appearing coast just as is

seen on the north and east sides of isolated mesas, or portions of the old

Plano Alto, which will be taken up in the following pages. Therefore,

I believe that the abruptness of the southeastern coast of Brazil is due to

its stability, as is shown by the absence of fossils, and to the southwest

dip of continental deposits, especially of late and post-Paleozoic age

which have been deeply eroded, and not to a post-Paleozoic faulting and

sinking of a land-mass into abysmal depths, for which no positive geo-

logical evidence exists. Much of the abruptness, however, is nothing

more than sea cliffs.

During Cretaceous times, there was a slight elevation of the coast of

central and northern Brazil. In comparatively recent times, the entire

Brazilian coast has risen 40 feet.1t This is sufficient to change the

earlier coast line and to produce another line which is abrupt, but due

to ages of erosion and not to faulting. Moreover, there are analogies

for this abrupt coastal condition in which no extensive sinking of a land- -

mass has occurred.

Therefore, summing up all the geological evidence, the lack of east

11 Hart, Krone and others have stated that the Brazilian coast has been elevated about

40 feet since the Pleistocene or Quaternary. I had ample opportunity to confirm this

view while traveling in Rio Ribeiro de Iguape and along the coast at various other

points of southeastern Brazil. The location and nature of the sambaquis (shell mounds),

the ocean caverns and wave marks along the bases of inland morros (hills) show that

the sea extended inland almost to the mouth of Rio Juquia (west of Iguape) even as late

as the time of the Indians in this region. In further evidence of this, marine shells are

dug up in wells east of Campos along the Rio Parahyba and along the west shore of

Lagoa Feia, which is now about eight miles from the seashore and about nine feet above

sea-level.

26 ANNALS NEW YORK ACADEMY OF SCIENCES

and west structural lines, the deep intervening sea, absence of islands,

relative evenness of great ocean depths, the absence of the deposits found

in deep seas on the continental shelves and the shape and abruptness of

the coast due to ages of erosion assisted by recent elevations, we conclude

that South America could hardly have been connected. with the eastern

hemisphere.*? )

PLANO ALTO AND THE PERMIAN INLAND BASIN

I have used the native term, Plano Alto, to include all of the sand-

capped tableland which extends south from the Guianas through Brazil

into Uruguay and west into Bolivia. The outlines of this region are

shown on Plate XII. All of these highlands appear to have been deposited

in a fresh-water inland basin during the Permian epoch. The remains

of Permian reptiles (Mesosaurus and Stereosternum), of the Gondwana

flora and other plants, of Scaphonyx, a Triassic reptile, of Schizodus,

Conocardium, Myalina and a few other marine lamellibranchs found in

the highland region indicate that the Plano Alto was deposited by wind

and rivers in a fresh-water inland basin of Permian age. The thin layer

of intercalated marine limestone indicates only a brief Permian in-

vasion of the sea in the Plano Alto of southeastern Brazil.

The Permian Inland Basin was almost surrounded by Archean moun-

tains: on the east by Serra do Mar and its northern spurs; on the south

and west by the Cordova Mountains and their southern spurs; on the

north by Archean rocks of Guiana and Venezuela, and on the north-

west perhaps by the Cordillera Oriental. The characteristic sandstone

found in all of this region was in part deposited in shallow fresh water

and in part shuffled about by winds into this Permian basin.

The basal Plano Alto formation or the floor of the Permian inland

basin is composed of granites, gneisses, crystalline schists and the like,

which are generally considered pre-Cambrian or Archean, because of

the absence of fossils. On various portions of this basal formation are

Paleozoic deposits which have already been mapped, but which will

without doubt be greatly extended as exploration proceeds. These maps

show that none of the Plano Alto included in Plate XII has been invaded

2 The absence of marine Lower Carboniferous fossils from eastern Brazil is not, in my

opinion, nearly as strange as the apparently entire absence of post-Paleozoic marine de-

posits of southeastern Brazil, in view of the fact that west of Serro do Mar are found

marine deposits of Devonian and Permian age. The first great escarpments in British

Guiana, not far from the coast, are due to the erosion of the Plano Alto sandstone in the

regions of the Kaieteur Falls; similar conditions exist right along the coast of south-

eastern Brazil. So it will take far better evidence than exists to prove that this coast

of Brazil is a post-Paleozoic faulted one and not a Paleozoic one which has remained

stable and has been changed by the erosion of later land sediments.

ANNALS N. Y. ACAD. SCI. VOLUME XXII, PLATE XII

Antafagartm

Blanco bncalain.

Puerta del Noarce \tu, fi

Voverer

Lor Andes

are!

Val per iaae

MAP OF PART OF SOUTH AMERICA

Showing the outline of the Plano Alto, which was deposited in the fresh-water Permian

Inland Basin and contains no Post-Paleozoic marine deposits

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 27

by post-Paleozoic seas. Hence continental and not marine deposits are

the more extensive deposits of Brazil.

The Permian inland basin is not uniformly symmetrical in reference

to the altitude of the basal rock in different sections, but its northern

and eastern sides are always higher than the southern and western.

There is also a considerable elevation of this basal formation between the

headwaters of Rio Paraguay (also seen in the iron and manganese de-

posits 600-800 meters above the level of Rio Paraguay in the Serra de

Urucum) and the Amazon, but this may be due to unequal early Paleo-

zoic erosion. This is especially true, if the streams flowed westward

before the Amazon was reversed. ‘This difference in elevation of the pre-

Cambrian and early Paleozoic or basal formation of the Plano Alto was

of the utmost importance for the formation of the Plano Alto. The

overlaps near these various higher (?) Archean elevations indicate that

much of the basal sandstone of the Plano Alto was derived from place

erosion of the higher points of the floor of the Plano Alto. Hence no

considerable extension to the east of present coast of South America was

necessary. The uppermost strata were deposited from the higher Archean

mountains on the east and northeast and hence dip toward the southwest.

The outlines of the Plano Alto shown on Plate XII may be slightly

extended by future work from southeastern Brazil toward Argentina, far-

ther into Paraguay and Bolivia, and farther west from Brazil toward the

Cordillera Oriental of Colombia and Ecuador. So much of the Plano

Alto has been deeply eroded and some entirely washed away that its exact

limits cannot be given at the present time.

Tt is evident from Plate XII that the Plano Alto proper is bounded by

parts of the basins of Rios Sao Francisco, Orinoco, Mamoré and various

other smaller rivers as well as by waterfalls and other changes in the

geological structure. In this connection, it is interesting to note that all

of the rivers which rise in the Plano Alto have clear water at least as long

as they flow on its formations, while many of the bounding rivers which

flow from the surrounding mountains often have yellow muddy water.

Rio Bermejo of Argentina, Rio Sao Francisco of Bahia, Rio Mamoré of

Bolivia, Rio Solomoes of Brazil, Rio Gurgueia of Piauhy and Rio Colo-

rado* of northern Patagonia are good examples of such rivers.

The yellow mud carried by all of these rivers, excepting Rio Sao Fran-

cisco, is produced for the most part by the erosion of Mesozoic and Ter-

tiary marine deposits; and inasmuch as these formations are not known

18 HWrom the size of the lower valley of this river and the identity of its fishes with

those of Rio San Juan, I am convinced that Rio Colorado was formerly much larger and

must have had some headwater from southwest Bolivia.

28 ANNALS NEW YORK ACADEMY OF SCIENCES

to exist in the Plano Alto, the yellow rivers assist in a most interesting

way to separate the Plano Alto from the rest of South America.

The original surface of the Plano Alto, as well as its uppermost strata,

dipped gently as a whole toward the southwest. This broad conclusion is

based on the following facts, after due allowance is made for ages of

erosion, reversal of rivers and the Tertiary rise of the Andes.'*

1. All of the streams and rivers which rise on the Plano Alto, 72. e., on

the Permian sandstone, at first flow south, southwest or west, even though

they afterwards flow north and east, 7. e., after they are eroded deeply

into the lower strata and flow over the older Paleozoic and Archean rocks.

For example, Rio Guaporé flows at first about 200 miles south and then

makes an elbow bend and flows west, north and lastly northwest into Rio

Mamoré. The streams of Jalapao in northern Goyaz, the headwaters of

Rios Parana and Paraguay and the streams of the Guiana highlands all

exhibit these same conditions in their headwaters. Even if stream piracy

is said to be responsible for these conditions, the general dip would still

be toward the southwest, because piracy could only be produced by the

more rapidly flowing northern and eastern streams robbing the head-

waters of the streams which flow toward the southwest, in order to explain

the existing conditions. ‘The cases of stream piracy considered in the

following pages show that it has been produced in exactly this way.

2. The second fact which supports the southwest dip of the Plano

Alto is that the north and east faces of isolated mesas or portions of the

original (not secondary) highlands are almost perpendicular, while the

south and west sides usually have gentle slopes. This is beautifully shown

by the Urucum Mountains near Corumba and by the many isolated mesas

in the Jalapao region of northern Goyaz. Also the west face of the Serra

de Parecis east of Villa de Matto Grosso is not nearly so perpendicular as

the east face of the Serra de Ricardo Franco, west of Villa de Matto

Grosso. The high Kaieteur Falls of an east-flowing river of British

Guiana and the Rio Branco flowing west from the same region having

only rapids support the same conclusion, 1. e., the surface of the original

Plano Alto dipped as a whole toward the southwest.

In view of all this, there appears to be no doubt that the Plano Alto

was previously much larger than generally considered. It has been very

stable since the Permian epoch. These facts may indicate a vast center of

evolution of plants and animals, but I hardly think so, for even at the

present time few plants and animals are able to thrive on this sandy ele-

vated region. In Permian and Mesozoic times, perhaps, this region was

14 The dip is so gentle that it is difficult to detect in the strata at exposed surfaces.

HASEMAN, GHOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 29

somewhat arid on account of higher, surrounding elder mountains, but

even if it had sufficient rainfall, on purely lithological grounds, a luxu-

riant vegetation and fauna would be impossible, because a sandy soil does

not retain the necessary constituents for a luxuriant growth of plants.

Besides, it appears more probable that unstable regions would produce

greater changes in living things than stable regions. Hence, not until the

Plano Alto was deeply eroded, could we expect to find a luxuriant growth

of vegetation and a complex fauna. Also, the Triassic Parana trap

which spread over much of the highlands of the states of Parana and Sao

Paulo of southern Brazil may have been an important factor in extermi-

nating and affecting the plants and animals, such as perhaps the Gond-

wana flora and Permian reptiles which are discussed in the second part of

this thesis.*®

EAST ANDEAN SEA

Several years ago, Professor Orton, Barrington Brown and others col-

lected marine or brackish water fossils mixed with fresh-water forms

along the Alto Rio Amazonas.

The following is a list of the fossils which have been ascribed to Alto

Rio Amazonas by Gabb, Conrad, Etheridge, Woodward and Boettger:

Anisothyris tenuis Gabb Hydrobia dubia Etheridge

carinata Conrad Lacuna (Hbora) crassilabra Conrad

obliquus Conrad (Nesis) bella Conrad

erectus Conrad Hemisinus sulcatus Conrad

cuneatus Conrad Melania tricarinata Etheridge

ovatus Conrad bicarinata Etheridge

hauxwelli Woodward scalaroides Etheridge

tumida Htheridge Dreissenia fragilis Boettger

amizonensis Gabb Turbonilla minuscula Gabb

Neritina (Isea) ortoni Gabb Corbula canamensis Etheridge

pupa Gabb Melanopsis browni Htheridge

puncta Etheridge Cerithium coronatum Etheridge

eiczac Htheridge Pseudolacuna macroptera Boettger

Hydrobia lintea Conrad Assiminea crassa Etheridge

confusa Boettger Bulimus linteus Conrad

tricarinata Boettger Anodonta batesi Woodward

(Dyria) gracilis Conrad

Fragments of the following genera have been reported by Woodward:

Myliobatus, Fenella, Thracia, Lutria, Anodon, Unio, Nautica, Odon-

stomia. Boettger has also reported Serpula (Vermes) and Rajidum and

15Jt is to be noted here that similar Permian and Triassic continental deposits exist

in other parts of South America (Ceara, Brazil, San Luis, Argentina, etc.) which are not

included in my Plano Alto, because they have been separated by marine deposits.

30 ANNALS NEW YORK ACADEMY OF SCIENCES

Percidarium (Pisces). Etheridge has reported parts of Chara (Plante).

Fossil tortoises and Mosasaurus also are said to have been found along

Rio Acre. These fossils are then a mixture of fresh-water, land, brackish

water and marine forms which lived apparently in a very special environ-

ment. ;

In reference to the age of the above fossils, I can do no better than refer

to Vol. XLIV of the Bulletin of the Museum of Comparative Zoology,

Harvard, pp. 25-27. In this résumé, Professor Branner states:

“Tf we grant that the upper Amazon region from Iquitos to Tabatinga is Ter-

tiary, there is no evidence that the mottled sediments of the lower Amazon are

of the same age, to say nothing of correlating them with similar looking beds

on the coast of Rio Grande do Norte, Parahyba, Pernambuco and Alagoas, 2500

miles away. This seems also to express Professor Derby’s view of the subject.”

Professor Branner also quotes Dall as saying that:

“The Pebas fossils are unique and difficult to determine the age because the

characteristic forms are extinct and have no obvious relatives. They may be

as old as Hocene or as young as Pliocene.”

The maps of the location cf the marine formations show that the entire

Plano Alto as herein mapped is Archean and Paleozoic; while the Andean

complex has an Archean nucleus more or less covered by marine deposits

of Mesozoic, and marine Tertiary is known to exist along both bases of

the Andes almost for their entire length. The trend lines of this region

are north and south, and they strongly indicate an extension of the Hast

Andean sea in the same directions.

The deposits along Alto Rio Amazonas are known from Pebas, Peru,

down Rio Solomoes as far as Sao Paulo de Olivenca, and south along Rio

Javary and Rio Acre. My maps show that this region lies between the

Cordillera Oriental of the northern Andes and the Plano Alto. It must

also be remembered that nowhere east of Sao Paulo de Olivenca, which is

1400 miles in a straight line from the mouth of the Amazon, have similar

or any post-Paleozoic marine deposits been found. This fact, as well as

others which have already been considered, strongly indicates that the sea

did not invade the Amazon Valley from the east, because the Plano Alto

would have been a permanent barrier to such an invasion from the Per-

mian to comparatively recent times.

The dip of the Plano Alto and the character of the sediment carried by

the Rio Negro, as well as the fact that no positive evidence exists which

warrants a northern extension of the East Andean Sea into Venezuela,

indicate a southern extension of the East Andean Sea, because as far as I

have been able to find, no Mesozoic and Tertiary are known to exist east

HASEMAN, GHOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 31

of the Cordillera Oriental of Ecuador or in the southern parts of either

Colombia or Venezuela, 7. e., as far south as the headwater of Rio Negro

in the Brazil-Guiana highlands. This vaguely indicates a connection be-

tween these Archean mountains of eastern Colombia (Cordillera Ori-

ental) and the Plano Alto, but inasmuch as this region is practically

unknown, too much emphasis must not be placed on such data.

There is then no positive evidence for either a northern or an eastern

extension of the Hast Andean Sea, but there is some positive evidence for

its southern extension. The location of the marine formations, the dip of

the surface of the Plano Alto and the character of the sediment carried

by southern Andean affluents of the Amazon vaguely indicate the same. —

The southern extension of Cretaceous deposits from the headwaters of

Rio Maranhao south toward Cochabamba is a good example of the south-

ern extension of part of this East Andean Sea which finally lost its con-

nection with the ocean on account of the Tertiary rise of the Andes.

In reference to the connection of this East Andean Sea with the ocean,

the following points are important :

1. Stelzner (1873) reported a sandstone containing marine bivalves

of (?) Tertiary age from Santa Maria, Catamarca of northern Argentina.

2. Brackenbush has reported marine or brackish water fossils of (?)

Cretaceous near the headwaters of Rio Bermejo which flows into the Gran

Chaco del Argentina. Melania was the most abundant form found, and

it may be that these deposits are fresh water and not marine.

3. Far to the west of the above regions, there is a great depression in

the Andean complex east of the bay of Arica and south of Lake Titicaca,

but so far there is no good evidence of a ys sea passing over the top

of the Andean complex.

4. It appears to be obvious from the characteristic fossils of the upper

Amazon Valley that a direct and broad connection with the ocean did not

exist. If, as the Cretaceous fossils of the Alto Rio Maranhao indicate, the

Hast Andean Sea existed during at least part of the Mesozoic to some

time in the middle or late Tertiary, there would have been ample time to

evolve this peculiar fauna of Pebas, because the conditions which would

have existed in this long, slender inland sea, into which many short rivers

carrying sediment flowed, would have been very different from those of

Patagonia, la Plata or along the Pacific slope.

It appears that the fossils of Alto Rio Amazonas are the last of the

fauna of the East Andean Sea which became buried in the mud carried

by the rivers into this vanishing sea, because the molluscs died with their

valves closed and because there are some fresh-water and land molluscs

mixed with the marine forms.

Beat ANNALS NEW YORK ACADEMY OF SCIENCES

In conclusion, then, we may say that the age of the fossils of the upper

Amazon Valley is not definitely known, and consequently we cannot more

than conjecture where the East Andean Sea joined the ocean ;*® but this

sea must have extended south and have had a narrow connection with the

ocean in the region of either Rio Bermejo, Rio Colorado-Patagonia or the

bay of Arica. This connection was perhaps not broken before the Mio-

cene, but it was broken by the Tertiary rise of the Andes. It is also im-

portant to note that no evidence is yet known which indicates that the

exit of this sea simultaneously cut east by west both the Archean moun-

tains flanking the northeast portion of Patagonia and those on the west-

_ern side of Patagonia in Chile in such a way as to isolate Patagonia from

Brazil. In fact, the evidence at hand is all against such a view.

REVERSAL OF RIO AMAZONAS

The following facts indicate that the direction of Rio Amazonas has

been reversed :

1. The Plano Alto which it now traverses has a general southwest dip.

2. The entire absence of marine Mesozoic and Tertiary fossils in the

Plano Alto.

3. ‘The position of the Hast Andean Sea.

4. 'The Tertiary rise of the Andes.

5. The general north and south direction of trend lines and the loca-

tion of marine deposits in western South America.

In an interesting treatise on the geology of the lower Amazon, Katzer

(1993) concluded that previous to the Miocene the Amazon flowed west

from somewhere in the region of Rio Paru. To the east and north of the

present mouth of the Amazon, he conceived a vast mass of land which

sank beneath the Atlantic Ocean when the Amazon became reversed.1*

Katzer also states that, during the reversal of the Amazon, a large lake

was formed, which extended eastward from Rio Nauta to the original

watershed. It appears that he has assumed the formation of this

huge lake in order to explain the formation of the fresh-water deposits

of Hreré. Katzer also maintains that no marine Cretaceous or Tertiary

beds exist in the lower Amazon Valley (Pirabas Cretaceous being on the

16 J am inclined to believe that it extended south along the east base of the Andes into

Patagonia, because there is no evidence of an eastern extension, a western over the

Andes, a northern into Venezuela. The connection by way of Gran Chaco is very ques-

tionable, for I believe the Plano Alto of eastern Bolivia joins the northern extensions of

the Cordova Mountains. Hence a southern extension into Patagonia meets no obvious

objection.

17 Pilsbry (1911) has also suggested the same idea in his studies on the distribution

of fresh water and land mollusea. It is interesting to note that three individuals work-

ing in different fields have quite independently arrived at the same general conclusion.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 33

coast) and that (?) Archean rocks underlie the alluvial deposits of the

Amazon and are continuous with the basal rocks of the highlands on

both sides of the Amazon.

The writer has independently, and from a slightly different stand-

point, arrived at the same general conclusion, namely, that the Amazon

is a reversed river, but he differs with Katzer in regard to the following

details :

1. The writer does not believe that there is any geological evidence

which even vaguely suggests either the existence or the submergence of

the hypothetical Jand-mass to the east and north of the present mouth of

the Amazon River. In fact, just south of the mouth of the Amazon

are marine Cretaceous fossils which indicate a slight elevation of the

present mouth of the Amazon from the sea and not a continuation to

the east with a hypothetical land-mass.

In this connection, no support for the hypothetical old land-mass can

be derived from the Barbados Islands, because they are entirely too far

to the north and belong to the Antillean complex. Both the absence of

islands and the deep-sea soundings are strong evidence that, previous to

the Miocene, no land existed east and northeast of the present mouth of

the Amazon across the Atlantic Ocean.

In fact, there is no necessity for the assumption of the supmergence

of any great mass of land to the east of the present mouth of the

Amazon, not even to form a watershed, because many rivers, like Rio

Paraliyba, Rio Tieté, Rio Iguassu, etc., of southeastern Brazil, rise within

a few miles of the Atlantic Ocean and then flow several hundred miles

before entering it.

2. The reversal of Rio Amazonas from the region of Rio Part is

roughly comparable to the supposed reversal of the Mississippi from the

region of New Orleans instead of nearer its headwaters. That is to say,

the comparatively recent reversal of the Amazon will have to be. I think,

in the region where the most of its large affluents enter it, namely, some-

where between Manaos and Santerem; because Rios Madeira, Solomoes

and Negro, which make the main stem of the Amazon, come together

just below Manaos. There is topographical evidence which shows that

the Rio Negro formerly entered the Rio Solomoes above its present

mouth, and hence the eastward extension of its mouth indicates that its

present mouth is near the region where the Amazon began to cut through

the old watershed, 7. e., near the dissected western base of the original

Plano Alto which has been washed away by the Amazon.

The arrangements of the low secondary sierras in the region of the

lower Rio Tapajos on the south side and near Obidos on the north side

34 ANNALS NEW YORK ACADEMY OF SCIENCES

of Rio Amazonas, taken along with the fact that the Amazon flows in

one channel only in two places (just below the mouth of Rio Madeira

and near Obidos), also strongly indicate that this is the region of the

previous watershed. Is it not to be expected that some trace of the

original watershed would still remain, if the reversal of the Amazon took

place during the Miocene? There certainly is not the slightest sug-

gestion of such a divide between Santerem and the mouth of the Amazon,

because the Amazon is so wide and swampy in all of this region, and

this is especially true in the region of Rio Part. Below Santerem, the

Amazon has many channels and many islands; between Santerem and

Manaos, there are two places where the Amazon flows in one channel, and

above Manaos Rio Solomoes (Amazon), there are again many channels

and many islands.

In view of all this, it appears that the original watershed must have

been somewhere between the mouths of Rios Madeira and Tapajos, and

I think that Obidos is near the actual point of reversal.

3. IT also do not believe that there is any evidence of a huge lake

which extended eastward from Rio Nauta to the old watershed. This

old view of Tertiary lakes has been ably combatted by Hatcher, Matthew

and ethers, but it will perhaps be worth while to consider the formation

of highland deposits in order that the fresh-water deposits of Hreré of

the lower Amazon as well as the other supposed evidence for a colossal

Amazonian lake will have an explanation. In the following brief con-

sideration, I have chosen the headwaters of Rio Guaporé, but those of

Rio Paraguay are equally as instructive.

Many streams, headwaters of Rio Guaporé, dash down the more or

less perpendicular faces of the so-called sierras from the flat surfaces of

the dissected Plano Alto into a large semicircular valley. This valley

includes the extensive Campos de Matto Grosso. Farther down this

valley, the campos are replaced by gigantic forests which encroach upon

the Guaporé so much that its channel is almost stopped up in several —

places below the Villa de Matto Grosso.

In this same region, the Serras de Ricardo Franco and de Parecis

(made by the Guaporé dissecting the Plano Alto) are much nearer to

the river than in the region of the semicircular valley. This serves as

@ block to the exit of the water which falls above this point. The dense

forest, fallen trees and water plants naturally assist in storing the heavy

rains which fall in the semicircular valley during the rainy season

(November to April).

In the middle course of the Guaporé, its channel widens very much.

Tremendous sand bars are encountered in each bend of the river. This

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 35

condition is replaced in the lower course of the river by a series of rapids.

after which it enters the Mamoré.

The Guaporé has then two great regions where sediments of different

nature are deposited at different altitudes and two great regions where

the water runs rapidly and carries away the eroded product.

Tt is the highland semicircular valley which, I believe, is roughly

comparable to the Serra de Ereré. This region is flooded yearly during

the rainy season, at which time much sand is deposited. This deposi-

tion of sand and other material (laterite, etc.) will eventually make a

secondary deposit of considerable extent, while the regression of the

headwaters of Rio Guaporé will eventually carry away all of the original

Plano Alto remaining in the surrounding Serras de Parecis, Ricardo

Franco and Agoaphey.

In fact, the Guaporé has already carried away more than one half of

the original highland formation from this region. The Rio Paraguay

is naturally assisting in this destruction of the “highest point.” As soon

as these two large rivers have obliterated the above mentioned sierras, the

semicircular highland valley will become one of the highest points (300

to 600 meters). After years of erosion on such a new high point, iso-

lated sections or mesas will be formed and secondary sierras like that of

Hreré will be produced which never were associated with colossal lakes.

I have observed near Corumba, Brazil, a similar deposition of leaves,

snails, ete., at two distinct levels. One of these levels is around the base

of the Serra de Urucum, including even deposits of limestone, and the

other is in the near-by pantanals (swamps) of Rio Paraguay. The same

process is going on near Sao Luis de Caceres, Brazil, and San Matias,

Bolivia.

Hence there is no evidence for a huge Amazonian lake and no neces-

sity for assuming one, for, as the writer conceives the reversal of Rio

Amazonas, it was a gradual process. The Tertiary rise of the Andes

did not suddenly close the exit of the East Andean Sea. but the water

cut deeply at the exit, until by the time this exit was almost closed,

stream piracy in the region of the old. divide (near Obidos) had pre-

pared a new exit for the water of the shallow East Andean Sea. For

some time, then, there were two exits for the East Andean Sea. As the

Andes rose higher and higher, the southwestern exit became closed and

all of the water rushed eastward. The amount of this water flowing

eastward was sufficient to wash away the old divide in a comparatively

short time.

Hyen the Rio Guaporé has washed away the most of the Permian for-

mation of the Plano Alto from a strip 50 to 150 miles wide. Is it not |

4—_NY

36 ANNALS NEW YORK ACADEMY OF SCIENCES

plausible, then, that the massive Amazon could have opened up its pres-

ent wide swampy valley and built up an extensive delta in a compara-

tively short time?

That the Amazon has swampy margins is no more an objection to its

reversal than the original swampy margins of the Mississippi would be

to its reversal. The Amazon, like the Mississippi, if it was reversed,

has washed away the most of the old divide.

Little or no exploration has been done far away from the forested

margins of the lower Amazon and its affluents. But an inland trip in

any of the region between Santerem and Obidos will usually reveal

sandy campos which are typical of the entire Plano Alto formations.

When this region is carefully explored, I feel sure that the old divide

will be definitely located.

The view which I have expressed concerning the Hast Andean Sea

offers a ready explanation of the origin of the peculiar marine-like fauna

found in Lake Titicaca. Lake Titicaca was doubtlessly once connected

with the East Andean Sea by a stream. When the Hast Andean Sea

began to disappear, some of its fauna entered or was cut off in the Titi-

caca basin. When the Andes rose still higher, the amount of rainfall

became more and more reduced until the exit of Lake Titicaca became

‘severed.

Higenmann'* states that Steinmann considers that the Titicaca basin

‘was a fresh-water basin whose southeastern exit was dammed by glaciers.

Glaciers may have assisted in closing the exit of Lake Titicaca, if they

were active at the time when the rainfall was so much reduced that the

precipitation scarcely exceeded evaporation; but it is evident that the

reduction of the rainfall due to the rise of the Andes was the more im-

portant factor. Otherwise, heavy rains would soon have filled up the

basin sufficiently to make a new exit. The Great Lakes of North America

were glaciated, and yet they made new exits. |

The absence of Manatus, Arapaima and Ostleoglossum, above the Ma-

deira Falls and their presence above the falls of Tocantins, Tapajos, and

in some of the coastal streams north and south of the Amazon are ex-

actly what one would expect to find according to all of the facts which

have been considered. ‘These three genera originally lived in the coastal

streams. When the Amazon became reversed, the Mamoré, which had

previously flowed southwest, changed its direction and suddenly formed

the Madeira Falls, which have been barriers to the migration of these

three genera.

Further zodlogical and topographical data could be given in support

48 Princeton Patagonian Reports, Pt. III, p. 372. 1909.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 37

of the reversal of Rio Amazonas, but inasmuch as the most of it is not

sufficiently decisive to account for such a momentous geological change,

it has been omitted; for such data may be said to harmonize equally well

with any view of the region in question.

Finally, in reference to the time of the reversal of Rio Amazonas, I

may restate that Katzer has placed it in the Miocene. It appears that

it could not have been before that epoch, but until the exact age of the

“Hast Andean fossils’ has been determined, we cannot settle this most

important question with any degree of certainty. The determination of

the exact age of the fossils from Alto Amazonas and the exact location

of the exit of the Hast Andean Sea offer alluring opportunities for future

exploration.

STREAM PIRACY

On the divides between various South American rivers, the headwater

streams sometimes approach one another with closest intervals. This is

particularly true of the divides on the Plano Alto. For example: Rios

Sao Marcos and Bartholmeo of Rio Parana; Rios Bezzero, Jardin and

Preto of Rio Paracatu and Rio Urucupa, both of which flow into Rio

Sao Francisco, and Rio Parana, an affluent of Rio Tocantins. All of

these rivers flow from brejos (swamps or sloughs), Lagoa Feia and

other highland lagoons between the villages of Paracattt and Formosa of

southeastern Goyaz. In fact, these headwaters of three large river basins

Tise in sight of each other from brejos which vary from 1100 to 1147

meters of altitude, and the maximum altitudes of the intervening sand

hills are never more than 1177 meters.’®

Stream piracy (coalescence of streams) may have existed on a very

limited scale between the following rivers—at any rate, their headwaters

are not far apart:

_ 1. Rio Ribeira de Iguape (? robbed Rio Capella) and Rio Paranapo-

nema.

®. Rios Sao Francisco and Doce and Rio Grande of the Parana near

Carandahy and Miguel Burnier, Minas Geraes.

3. Rio Parahyba and Rio Tieté, Rio Parahyba having robbed its

headwaters flowing south from Rio Tieté.

4. Rios Araguay, Xingu and Tapajos and Rio Paraguay.

5. Rio Ibicuhy of Rio Uruguay and Rio Vaccachy of Rio Grande do

Sul.

19 The highland fauna and flora certainly interchange between these three river basins,

but the typical Amazonian, Sao Franciscan and Paranean fauna and flora are at least

one hundred miles away from this headwater region, 7. e., at a much lower altitude.

38 ANNALS NEW YORK ACADEMY OF SCIENCES

6. Rio Sao Francisco and Rio Itapicurt east of Joazeiro.

?. Rio Agua Branca and Rio Negro of Rio Sao Francisco and Rio

Palma of Rio Tocantins.

8. Rio Grande of Rio Mamoré and Rio Pilcomayo of Rio Paraguay.

9. Rio Branco of Rio Negro, Rio Part, Rio Trombetas and Rio Esse-

quibo.

10. Corrego da Boa Ventura of Rio Guaporé and Corrego de la For-

tuna of Rio Paraguay.

Even if these streams, however, were previously connected, the con-

nection would have been so small and at such altitudes that nothing

but the highland fauna could have interchanged. JBesides, all these

streams have and have always had waterfalls in some part of their

courses. |

Plate XIII shows in a general way that no continuous waterway exists

between Rio Paraguay and Rio Guaporé, as has been so often erroneously

stated. Several years ago, an attempt was made to cut a canal between Rio

Guaporé and Rio Jaurt, but it was given up on account of the intervening

clistance (about 20 miles) and the nature of the material to be removed.

c. grandson of the man who attempted to make this canal hauls rubbe2

over this divide in an ox cart, but it is far more difficult than by the

Bolivian trail from San Ignacio past San Matias to Descalvados.

The former trail (Villa de Matto Grosso by the way of Jaurt to Sao

Luiz de Caceres) is only 301 kilometers long, while the latter trail ( Villa

de Matto Grosso to Bastos, las Encruzijadas, San Matias and Descalvados

or Sao Luiz de Caceres) is 488 kilometers, but it is much smoother.

While following this latter trail, I had ample opportunity to observe that

no connection exists between the Rios Alegrete and Agoaphey. In fact,

Rio Santa Rita flowing off this same sierra has one waterfall about 400

feet high. All of these rivers are nothing more than creeks.

The writer found that the nearest as well as the lowest approach be-

tween the headwaters of Rios Guaporé and Paraguay is in the region of

the Corrego de Boa Ventura and la Fortuna; but even though there is a

break in the continuity of the Plano Alto in this region, firm hills of con-

siderable height separate these creeks, which are not more than ten feet

wide and three feet deep and at least four miles apart.

Dr. Alipo Miranda de Ribeiro, Secretary of the Brazilian National Mu-

seum, was a member of the telegraph commission which has explored the

northern portion of the Paraguay River. He explored Rio Jaurti and

Sepatuba and found large waterfalls in each of these rivers. On the other

side of the divide, the commission found two large waterfalls in Rio

Juruena which flows into Rio Tapajos. One of these falls was about 400

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feet high. Similar conditions are said to exist between the headwaters of

Rios Paraguay, Tapajos and Xingu, all of which rise on the Plano Alto

and are not connected.

Stream piracy on a large scale has taken place between Rio Sao Fran-

cisco and Rio Tocantins and between Rio Negro and Rio Orinoco.

Rio Orinoco is robbing Rio Negro of one of its previous affluents, the

Canal de Cassiquiare. This is due to the fact that there is a long gentle

slope of the highlands toward the Amazon Valley, while the northern and

eastern slopes are more abrupt. On account of these conditions, Rio

Orinoco is still cutting rapidly and deeply through a so-called sierra just

above its connection with the Canal de Cassiquiare. The Rio Negro on

the other hand is not cutting as rapidly in this region as Rio Orinoco.

The Canal de Cassiquiare is a long, narrow, lakelike mass of water with

high sand hills along and near either bank. The Rio Negro end of this

“canal” is being gradually stopped up by secondary deposits of sand and

plants, and consequently as soon as Rio Orinoco cuts more deeply into the

hard basal highland rock, it will drain the Canal de Cassiquiare and

appropriate it as an affluent.

Lakes and swamps similar to the Canal de Cassiquiare are quite com-

mon in and on all of the Plano Alto. Sete Lagoas, at the headwaters of

Rio Paraguay, and Lagoa Feia, near Paracatu, in southeastern Goyaz, are

examples of such lakes or swamps; but inasmuch as no marine deposits

are known to exist in any of these regions, it appears that all such lakes

are entirely due to unequal erosion and changes in the courses of the high-

land streams. Such is evidently the case of Canal de Cassiquiare, for it is

almost surrounded by the highland sandstone, and in the regions of the

upper waterfalls both Rio Orinoco and Rio Negro flow over the hard

ancient basal highland rock. The rocks in the Rio Negro Falls are essen-

tially the same as those of the Madeira Falls.

Similar conditions exist between the headwaters of Rio Tocantins and

Rio Sao Francisco in the Jalapao portion of northern Goyaz. Wells

(1875) and Froh (1907) reported a connection between these rivers; but

my discovery, which was entirely independent, was based on the presence

of certain Amazonian fishes not previously reported from the Sao Fran-

cisco River.?° .

Plate XIV shows that an affluent of Rio Tocantins and one of Rio Sao

Francisco rise in the Brejo de Varedao, which is surrounded on all sides

*0Tt may be of some interest to note that I do not now consider that the presence of

these fishes is evidence for this connection, because I later found the same fishes in other

rivers where no connections exist. My erroneous assumption, however, which is a com-

mon one among static zodgeographers, led me to the discovery of several facts.

40 ANNALS NEW: YORK ACADEMY OF SCIENCES

by massive sections (1. e., faces of mesas, etc.) of the Plano Alto. These

sections appear like many flat-topped sierras, but are nothing more than

the perpendicular faces of the remains of a dissected plateau. From a

distance, these sections look like gigantic modern fortresses; but as one

approaches nearer and nearer to them, he notes that there are immense

sand hills (campinas), which slope to and away from these sections of the

Plano Alto. That is to say, the bases of the perpendicular faces of the

sections are about 1000 feet below the top, while the tops of the campinas

which join the same bases are often not more than 500 feet lower than

the top of the sections. Hence the result of erosion is very striking in all

the region of the Plano Alto, since the original high points tend to be-

come the lowest.

The explorer is impressed by the great number of the highland brejos

which flow along parallel lines and are about 6600 meters apart. These

brejos are often less than 100 feet wide and usually have a small, rapid,

clear stream in the center. The edges are so boggy and full of palms and

other typical swamp plants that one often encounters great difficulty

either in getting water to drink or in crossing them.

The Brejo de Varedao appears to be due to unequal erosion and a con-

sequent deposition of sediment about the ends of the swamps. This region

offers a peculiar case of stream piracy unless Rio Sapao has cut back into

the headwaters of both Rio Nova and Rio Formosa. At any rate, the

Brejo de Varedao is gradually being drained by Rio Sapao, which is ecut-

ting through the hard basal highland rock in its rapid descent to Rio Sao

Francisco on one hand and by a similar action of Rio Formosa, which

flows out of the opposite end of the same swamp to Rio Tocantins. Dur-

ing heavy rains, the Rio Nova still assists in draining this swamp. In

fact, it is almost impossible to ride between its headwaters and the edge

of the Brejo de Varedao, except during the dry season.

It was with great difficulty that the writer pushed his way through the

swampy margins of the Brejo de Varedao to its lakelike central part,

which could not be fathomed by the longest poles obtainable. In the

middle, there was no current, but at the opposing ends were the rapid Rio

Sapao and Rio Formosa, which at once leave the swamp, flowing over the

basal highland formation. These streams are on the average 20 feet wide

and four feet deep during the rainy season. At this time the Brejo de

Varedao is about eight miles long and less than one mile wide at its

widest part.

During the rainy season, a canoe can descend from the Brejo de Vare-

dao with little difficulty as far as the great cataracts of Paulo Affonso of

the lower Rio Sao Francisco; but a canoe cannot descend so easily from

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 41

the Brejo de Varedao to Rio Tocantins, on account of a waterfall in the

lower course of Rio Nova, known as Cachoeira da Velha.?+

The Cachoeira da Velha is about 200 feet wide. It had, when I saw

it, a small island with two trees. These trees are evidence that the chan-

nel was originally a few feet to the north of its present site, where there

is still evidence of an old side channel which had no perpendicular falls.

During half-flood season, I estimated the highest point of the falls at 24

feet. The north side was 12 feet and the south side was a rolling mass of

water about six feet high. The south side has nearly all of the water,

because there is an abrupt bend in the river about 500 feet above the falls.

‘These falls are not any higher than the Theothono Falls of Rio Madeira,

which have been passed by nearly all of the Amazonian fishes. It must,

however, be remembered that Rio Madeira has a volume almost as large

as the Mississippi and that the Theothono Falls are only about 100 meters

above sea level, while the Cachoeira da Velha has a small volume of water

and is‘at least 300 meters above the sea level.

The rocks of the waterfall are the same general type of (?) pre-Cam-

brian found in various waterfalls of the Plano Alto region. I picked up a

piece of flint which I believe corresponds to the Jacuipe flint of Serra de

Jacobina. To the northeast of this region, the iron, manganese and dia-

mond-bearing gravels were observed near Paranagua, Piauhy. This indi-

cates a similar structure as the Jacobina series of Bahia.

The dissected faces of the Plano Alto are far away from the south side

of the Cachoeira da Velha and the river does not flow off of a mountain,

as sketched by Froh. On the north side of the Cachoeira da Velha, there

are several mesas of the dissected Plano Alto between Rio do Somno and

Rio Nova.”?

BARRIERS TO AQUATIC MIGRATION AND CONDITIONS OF ENVIRONMENT

Under the above heading, I will not consider physiological barriers, i. e...

if a fish is found in the upper course of a large river like Rio Paraguay,

which is navigable by small ocean steamers, and is not found in the lower

course of the same basin (Rio Parana), we may look on this restricted

distribution as being due at least in part to a physiological barrier.

Under the subject of barriers to aquatic migration, I will discuss only

one of the geological barriers, namely, waterfalls. Too much’ emphasis.

cannot be placed on the location, size and character of the numerous

It is so named because an old fish-like woman is supposed to have drowned a fleeing

bad Indian at this point about 100 years ago. The natives are afraid to go to this

waterfall.

22 Canoes descend from Porto Franco on Rio do Somno to Para during the rainy season.

42 - ANNALS NEW YORK ACADEMY OF SCIENCES

waterfalls, because they mark changes in geological structure which are

associated with profound changes in the environment of both aquatic and

terrestrial life. The waterfalls are also very important, because they have

been the source of some errors made by previous writers on the distribu-

tion of the aquatic forms. ‘hese errors have for the most part been due

to a lack of exact data concerning the waterfalls.

The first of these great waterfalls is Paulo Affonso, which may be

designated as the “King of Cataracts.” Paulo Affonso is found in the

lower course of Rio Sao Francisco and consists of a great series of rapids

above and below the falls. The volume of water which rushes over Paulo

its level 200 meters between Jatoba and Piranhas, a distance of 121 kilo-.

meters by the railroad which connects the navigable portions of the river

above and below the real falls. In fact, the Rio Sao Francisco has lowered

Affonso is on the average about the same as that of Niagara. Nearly ali

this “muddy” water comes from the precipitation in the headwaters of

Rio Sao Francisco, its lower course from Joazeiro to Piranhas seldom

having rain. Consequently, during the dry season, Paulo Affonso is much

higher and more beautiful because the water is clearer. The relative

difference in the height of Paulo Affonso during the dry and flood seasons

is very great, because there are seven principal channels in the wide river

above the falls through which the water rushes into the long, narrow gorge

below the falls. I estimated the maximum height of a series of tumbles

at 143 feet, but the flood-marks in the gorge below the falls for the pre-

ceding year, which had only an average flood, would have reduced this

height at that time about 40 feet.

Of the seven principal channels at Paulo Affonso, the southernmost one

on the Bahian side leaves the river about two miles above the falls and

gradually rushes into the gorge below the falls. The natives claim that

fishes swim up this channel during large floods, and I have no reason to

doubt their statement, because I found that the fishes above and below

the falls were almost identical, and because no great perpendicular fall

exists in the southernmost channel of Paulo Affonso. Naturally the ma-

rine forms which enter the mouth of this river are not found above the

falls, but Pachyurus (Corvina) is; and since it is never found in either

Rio Novo or Rio Sapao, it must have originally passed Paulo Affonso,

when it left the sea and became a permanent fresh-water form.

Finally, I may state that Professor Branner and one of his students,

Mr. R. Crandall, have recently reported marine Cretaceous above Paulo

Affonso at Jatoba. Consequently, Paulo Affonso did not exist at that

_ time, but most of the present fishes of the Sa Francisco also did. not

exist at that epoch. There can be no doubt that Paulo Affonso is older

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 43

than nearly all if not all of the fishes which are found in the Rio Sao

Francisco. Therefore, Paulo Affonso appears not to have been a barrier

to the migration of fishes.

In violent contrast to Paulo Affonso are the Iguassti Falls of Rio

Tguasst. I say violent contrast, because the Rio Iguassu flows from the

sandy highlands of Parana, Brazil, and after rushing over the Parana

trap rock at several lesser waterfalls, finally tumbles over the Iguassu

Falls and flows calmly toward the sea. The Iguasst Falls are about 200

feet high, and even though they are not exactly perpendicular, there are

no side channels which could have been passed by fishes. The average

volume of water is about one third more than that of Niagara Falls.

The Iguasst. Falls tumble over the “Parana trap,” which has been de-

termined as Triassic by Professor Derby, and they are so old that they

appear to have been an absolute barrier to the migration of the fishes.

Above the falls, I found only about 25 species of small, common highland

genera, many of which are slightly different from the same species in the

neighboring highland streams. These differences, for all we know, may

or may not be inherited, because the external conditions in the Rio

Iguassu basin are different from those in the neighboring rivers. At any

rate, the Iguasst Falls have been a permanent barrier to the migration of

lowland fishes and the deplorably poor ichthyological fauna has for the

most part evolved from the common highland fauna, which will be con-

sidered in the last part of this paper.

The next great South American waterfall which will be briefly con-

sidered is that of “Sete Quedras” (Brazilian) or “Guayra” (Guarani).

The Guayra Falls are in the middle course of Rio Parana, between the

State of Parana, Brazil, and the Republic of Paraguay. The Parana

River above the falls is 4200 meters wide, and the water rushes through

seven channels over seven falls of varying height and width. The water

then unites in a narrow gorge, which is only 80 meters wide, and rushes

madly, with a deafening roar, for about 40 miles before its force is spent,

and then Rio Parana calmly rolls on toward the sea. A small Argentine

man-of-war went up the Parana as far as the mouth of the Iguasst River,

which is several miles below the Guayra Falls, and sailed up the Iguassti

River as far as the Iguassti Falls. Hence these rivers are not as impass-

able as many are inclined to believe. At any rate, fishes can and have

passed the Guayra Falls, because I found that the ichthyological fauna

which lives in such rivers as the Alto Parana and its affluents is identical

above and below the falls.

The Pirapora Falls of the Alto Rio Sao Francisco have been errone-

ously stated to be a barrier to the migration of fishes. In fact, the Pira-

44 ANNALS NEW YORK ACADEMY OF SCIENCES

pora Falls are not really falls at all, because during the rainy season they

entirely disappear. I fished in their edge during the beginning of the

rainy season and found them no barrier to the migration of fishes.

It is interesting to note that “Pirapora” has been derived by some from

pira = fish and pora= port, but I found in Paraguay that the Guaranis

spell pora as pona (= beautiful), but pronounce it as pora; hence the

word would mean “beautiful fishes” and not “fish port.” Hither of these

names might lead one to think that fishes do not pass these falls, but dur-

ing the dry season Pirapora becomes a veritable fish port, because the

fishes wait below the falls for the rise of the river at the beginning of the

rainy season in order to go farther up the river to spawn. This move-

ment upstream is well expressed by the Guaranis by the word piracema

(pira = fish, cema = rising).

Another interesting waterfall is that of Piracicaba, found in Rio Piraci-

caba in the State of Sao Paulo. This fall has been a barrier to most

species of fishes, but it has not been the absolute barrier that its name

indicates (pira = fish, cicaba==ends). During Piracema the Indians,

in their exaggerating way, say that the fishes pile up so deep below the

waterfalls distant from civilization that one can walk across the rivers

on the backs of the big fishes, and that their wriggling about produces a

roar like thunder. Even though this is extremely exaggerated, there is a

certain element of truth in it.

In order not to swamp the reader with too much detail, I will directly

draw my general conclusion concerning waterfalls, because I have already

described the important types of fall and because the remainder of the

falls have been described with varying degrees of accuracy by numerous

other travelers and explorers. Before drawing these conclusions, it is

worth noting that whenever the height of a waterfall is given, it is the

maximum distance between the water levels above and below the falls,

and not the height of side channels and series of “tumbles” which often

furnish ready passages for fishes and other aquatic forms.

Practically none of the many hundred waterfalls existing in the cen-

tral and lower courses of Rio Parana and its affluents. Rios Tieté. Grande,

Paranaponema, Ivahy and Tibagy, Rio Uruguay, Rio Tocantins, Rio

Xingti, Rio Tapajos, Rio Madeira, Rio Negro and other South American

rivers have been barriers to the migration of the fishes, which either

spawn in the upper courses of the rivers or live in such environments.

The only absolute exception to this conclusion is the Iguasst Falls, which

have already been considered.

The Giral, Theothono, Guaja Mirim and the other 27 Madeira-Ma-

moré falls, Esperanca of Rio Beni, Alcoboca and numerous other falls

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 45

of Rio Tocantins, Forto Principe da Beira of Rio Guaporé, Avanhandava

and Itapura of Rio Tieté, Salto Grande da Paranaponema of Rio Parana-

ponema, Urubupunga and Dorado of Alto Rio Parana, Itaituba of Rio

Tapajos, ete., are typical examples of such falls, the general location of

which are shown on Bartholomew's New Commercial Map of South

America.

In the headwaters of practically all of the highland rivers flowing

into either the Amazon or the La Plata, there exist and have existed in

some part of their courses many very high waterfalls which are now, and

their ancient locations far away from their present sites have been since

the early Mesozoic epoch, effective barriers to the migration of all aquatic

forms, excepting the common highland fauna, which is always very poor

in species. These fails are even now, after ages of erosion, often 400 or

more feet high and are nearly always perpendicular. ‘Their volume of

water is small and no possible side channels exist. The Kaieteur Falls

in Guiana, Salto de Rio dos Patos in Parana, Brazil, the fall in Rio

Santa Rita from Serra da Agoaphey and the fall in Rio Juruena of Rio

Tapajos are typical examples of such falls.

Between the waterfalls of the central and lower courses of the rivers

and the waterfalls in their headwaters, as well as below the waterfalls in

the central and lower courses of the rivers and along the Atlantic coast

of South America, are many extensive swamps which have been produced

in various ways. It is these swamps which are responsible for a common

erroneous opinion that the most of South America consists of swampy

sultry lowlands which are beladen with every bad thing that exists.

Even though these swamps are not as extensive as they are generally

considered to be, they are nevertheless of profound significance, espe-

cially in the study of environment and the distribution of aquatic forms,

and therefore we will next consider the mode of origin of these swamps,

which may be roughly divided into two classes, as follows:

1. The coastal swamps, which have been formed by the comparatively

recent slight rise of the coast, followed by an unequal deposition of sedi-

ment both by the rivers and by the production of sand-dunes. ‘This is

true for the Lagoa dos Patos of Rio Grande do Sul, as well as for the

various other coastal swamps like those of Iguape and at the mouth of

Rio Doce. There appears to be little or no doubt that the swamps of at

least part of the Gran Chaco of Argentina and southwestern Paraguay, as

well as those at the mouth of the Amazon and Orinoco, have had a similar

origin.

2. In contrast to the above type of swamps are those along the central

and upper courses of Rio Paraguay, known as Pantanals, and Rio Ama-

zonas and all of its highland affluents, as well as all the swamps along the

46 ANNALS NEW YORK ACADEMY OF SCIENCES

bases of the remains of the Plano Alto. These swamps are entirely due

to unequal erosion and a consequent deposition of sediment at different

levels, and this has been assisted by the growth of a dense flora and

changes in the channels of the rivers. The Concepcion Lake of Bolivia

and the Uberava Lake of Rio Paraguay and a countless number of other

lakes which are found within the limits of the Plano Alto (cf. Plate XII)

belong to this class of swamps. The swamps in the central courses of the

rivers are usually due to changes in their channels.

During the rainy season, all of these swamps become flooded, and con-

sequently it takes about a month of heavy rains to start a large flood in

the lower courses of the rivers. It also takes three or four months of the

dry seasons before the water can percolate through the swampy vegetation

and get into the channels of the rivers.

The location and heights of the waterfalls, the location and kind of

swamps and the kinds of environment are directly associated with the

_ geological structure and the altitudes. Inasmuch as I have already briefly

considered the geology, the altitudes may be profitably considered at this

point.

The highest point in Brazil is Pico de Itatiaya (2804 meters), which is

between the states of Rio de Janeiro and Minas Geraes. The highest

point in Guiana is 2621 meters. The highest point in the Cordoba Sierra

is 2530 meters. A few peaks of the Andes surpass 20,000 feet, and Acon-

cagua is over 22,000 feet high. The Plano Alto is seldom over 1000

meters, but one point in Goyaz is said to be 1500 meters above sea level.

In Rio Grande do Sul,. parts of the eroded Plano Alto are often not more

than 300 meters above sea level.

_ The divides formed by these higher elevations are, as has been previ-

ously stated, correlated with the natural conditions of the descending

rivers. Therefore, the following brief list of altitudes along the more

important rivers has been given, not only on account of the above reason,

but also because the abundance and the distribution of all plants and

animals are confined between more or less definite positive and negative

altitudes of the land and the sea.

Place above seuieeet

Sao Antonnio, last fall in Rio Madeira.......................... 96 meters:

Guaja Mirim, first large fall in Rio Mamoré..................... 203s

Jatoba, above the Paulo Affonso Falls........................0- 246 “*

Altocdo: Serra; near Sao: Palo. a secrmeins oe ees ane on ee 837 sf

Rio WMeté, near Sao Paulo............-...-+.5-+ 22+ 52s. -about.. 0s

Jalapaio, headwaters of Rio Sapao....................... about.. G00 ‘

Mouth of Rio Jaura in Rio Paraguay.......................-.- 153 ose

Sao Carlos, on Rio Negro above Manaos..............---ee+-eeee ZAT —s

Villa de Matto-Grosso, on Rio Guaporé................... about.. 300 “

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 47

The above brief list of altitudes is sufficient to show

1. That the upper courses of the highland rivers are at least 150 meters

and usually more than 200 meters above sea level. In the upper courses,

I do not include the headwaters which naturally correspond more or less

with the altitudes of the divides. In this list, it is important to note that

Rio Paraguay has about the same altitudes as Rio Amazonas and stands

in marked contrast with that of the Alto Rio Parana and its affluents like

Rio Tieteé.

2. That the headwaters are rarely less than 300 meters and on the

average are about 900 meters above sea level.

3. That the middle and lower courses of the rivers are usually less than

200 meters and on an average are 100 meters above the sea level. The

main stem of the Amazon and La Plata rivers are exceptions to this rule,

because their middle courses are only about 100 feet above sea level.

The marked differences in altitude, which are more or less associated

with the direction of the wind, the temperature, the amount and time of

rainfall, the geological structure and all the other factors which compose

an environment, separate South America into several distinct faunal and

floral regions, even though most of it is generally considered to be tropical.

Space will not permit a detailed consideration of the composition of

faunal regions, but perhaps the following rough analogy will make the

idea clear. When the naturalist who is familiar with the United States

thinks of the regions near Tampa, New Orleans, New York, Chicago,

Flagstaff and San Francisco, he can easily distinguish any one of the

regions from the others in various ways. In like manner, the naturalist

who is familar with South America can at once distinguish any one of

the following regions from the others: The Pampas, the Campos, the

Plano Alto, Patagonia, Gran Chacos, Mattos Grossos, the Secca or arid

regions of Brazil and the Cordillera de los Andes.

In brief, when I think of Patagonia, I think of a region covered with

calafate bushes, low, level or rolling, scanty rainfall, temperate climate,

and a region characterized by a general paucity of life, but not a desert.

When I think of Pampas, I think of a deep, rich alluvial soil, whose

grassy surface is as level as a floor and in many ways is like an Illinois

prairie which never freezes. When I think of the Plano Alto and Campos,

I think of more or less elevated sandy plains which are covered with

scanty highland grasses, an occasional scrubby tree, sparkling water, few

pests and a delightful climate. When I think of Chacos, I think of

swamp palms, Lepidosiren, cutting grasses, floating treacherous grassy

surfaces, decaying plants, foul odors, sultry atmosphere and alligators.

“When I think of the yast Secca country of northeastern Brazil, I think

5—NY

48 ANNALS NEW YORK ACADEMY OF SCIENCES

of little or no rainfall, cactus, spiny and thorny plants and scrubby trees,

dead insects, starving animals, hot dry air, and a gray dead appearance of

everything excepting the sun, moon and stars.

When I think of Mattos Grosses, I think of forests and jungles which

are composed of an amazing number of kinds of plants all mixed up,

loaded with vines and orchids and harboring monkeys, parrots, sloths,

armadillos, humming birds, insects and other animals galore. In other

words, a region where life is at its present climax, a region where the con-

ditions for the existence and evolution of life are most favorable.

In like manner, we can arrange the rivers of South America into a few

groups in which the sum total of the natural conditions are almost iden-

tical, excepting the volume of water. These groups of environmental

complexes are shown on Plate XV and are as follows:

1. In the Campos of the State of Rio Grande do Sul and Uruguay is

Rio Uruguay on one side and Rio Grande do Sul on the other side of the

divide.

2. Rio Parana and its affluents, Rio Tieté, Rio Grande and Rio Parana-

ponema with their cool, clear water rushing over many waterfalls, are on

the west slope of Serra do Mar, and Rio Ribeira de Iguape, Rio Parahyba

and Rio Doce on the east side of the same sierra.

3. The northern part of Rio Sao Francisco belongs to the “secca” re-

gion, which includes the most of Rio Parahyba do Norte; the western part

belongs to the highland region, which is also embraced by the Amazon,

and the southern part of Rio Sao Francisco belongs to the region of Alto

Rio Parana.

4. The Amazon is so diverse that it includes several smaller regions ;

and Rio Essequibo and perhaps Rio Orinoco on one side and the Alto Rio

Paraguay on the other side belong to the Amazonian realm in reference

to the sum total of their natural conditions.

5. The Patagonian and the West Andean rivers each belong to quite

distinct natural regions. Perhaps the Patagonian should include the

southern portion of the West Andean, 7. e., Chile.

The profound importance of a clear conception of these regions will, I

hope, become more evident in the last part of this thesis.?°

22 Tt is well known that the environment produces changes in both the individual and

the species. When these changes are produced in the germplasm, they are transmitted to

the offspring. Is it not to be expected then that at least some of the offspring from the

same species of plants and animals would become different after the parent species ar-

rived in the different environments? And may it not also be expected that at least

some of the offspring from the same species under identical external conditions, even if

widely separated, will evolve some identical species?

ANNALS N. Y. ACAbD. Sct. VOLUME XXII, PLatE XV

OUTLINE MAP OF SOUTH AMERICA

Showing environmental complexes in which the sum total of the natural conditions are

about equal

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 49

Part Il. DistrRiBuTION OF THE SouTH AMERICAN FISHES AND ITS

BEARING UPON ALLEGED CONNECTIONS BETWEEN SOUTH

AMERICA AND THE HASTERN HEMISPHERE

DISTRIBUTION OF THE FISHES

Introduction

In attempting to explain the distribution of South American fishes, I

haye been handicapped by lack of data bearing upon

(a) The question whether all of the species concerned are real species.

Experimental work is entirely lacking which would demonstrate whether

the species are real, composed of several elementary species, varieties or

only individual and local variations (ontogenetic species of Jordan)

which have responded to different external conditions, 1. ¢., affecting the

somaplasm and not the germplasm.

(b) The nature of the ancestral species. That is to say, which species

of a given genus is the nearest to the ancestral one that became widely

distributed.

mentary.

If I followed the old static method, not only such data but also the

changes wrought by the environment would almost be neglected. In this

case, it would be relatively easy to compile the exact localities of all of

the species in a given family, and by comparing the long list so obtained,

divide the world into as many faunal regions as these data would warrant:

In strong contrast to this older method used by the most of the writers

on geographical distribution of South American animals is the study on

Leptinotarsa by 'Tower.** I cannot improve on his statement (p. 52)

concerning the two viewpoints of geographical distribution, which is as

follows:

“The geographical distribution of animals, or animal geography, is usually

considered from one of two viewpoints, the static or the dynamic. Considered

from the static standpoint, the facts of distribution are taken and arranged

according to some empirically chosen standard, and zones, Sub-zones or other

unnatural areas of distribution are established. The study of animal distribu-

tion from this standpoint is a dead and profitless pursuit. Dynamically con-

sidered, animal geography seeks to explain the facts of animal distribution as

we now find them in terms of the relation of the animals to each other and to

their environmental complexes.”’ ;

24 W. i. Townr: “An Investigation of Evolution in Chrysomelid Beetles of the Genus

Leptinotarsa.” Carnegie Inst. of Washington. 1906.

50 ANNALS NEW YORK ACADEMY OF SCIENCES

In so far as possible, it is the dynamic aspect of the distribution of the

South American fishes that I shall consider in the following pages.

On page 9 of the same remarkable book, Tower states the following ten

criteria which have been used by Adams, Allen and others to determine

the center of origin of a fauna when no fossils are known:

1. Location of the greatest differentiation of a type.

. Location of dominance or greatest abundance of individuals.

. Location of synthetic or closely related forms.

. Location of maximum size of individuals.

. Location of greatest stability and productiveness in crops.

. Continuity and convergence of lines of dispersal.

. Location of least dependence upon a restricted habitat.

8. Continuity and directiveness of individual variation or modifica-

tions radiating from the center of origin along the highways of dispersal.

9. Direction indicated by biogeographical affinities.

10. Direction indicated by animal migration in birds.

After critically considering the above ten criteria, Tower?’ states that

the following four are adequate for determining the centers of origin or

adaptive radiation, without the introduction of any of doubtful value:

1. Location of greatest differentiation of a type.

2. Continuity and convergence of lines of dispersal.

3. Location of synthetic or closely related forms.

4. In some cases, location of dominance or great abundance of indi-

viduals.

I quite agree with Tower that these criteria are sufficient to determine

the point of origin, if the four criteria themselves can be correctly deter-

mined, and I believe that these four criteria can be correctly determined

in a given genus like Leptinotarsa, but when the genus becomes very

widely distributed, with many distinct species and varieties, it is more

difficult, if not impossible, to determine these four criteria. At any rate,

they cannot be correctly determined for the families and orders of animals

without the aid of fossils, because the factors and conditions become so

complicated that the determination would be nothing more than an

opinion of the individual. For example, I have been able to show that

certain species in a certain genus of Cichlide have evolved from another

species, but which of the many species in a widely distributed genus is

the ancestral one cannot be easily and correctly determined. Further-

more, even though we are able to determine that one particular genus of

the twenty odd genera of the cichlids is the most primitive and that most

~2 S> Ot H CO

2 Op. cit., p. 13.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 51

of the genera are found in the Amazon valley, we have yet absolutely no

evidence that the ancestral Cichlide originated in the Amazon. In fact,

as Matthew has already pointed out, the presence of a large number of

species in a given locality is no evidence as to their point of family origin.

Indeed, neither the genus nor the family need have originated there, since,

as Matthew has shown, the point of origin is apt to be the first place of

extinction. We may note, therefore, that the above rules which have been

used to determine the point of origin of any group of animals have only a

hmited application.

So far, there have been only a few specialists interested in the ichthy-

ology of South America. Of these, Professor EHigenmann has certainly

given us the best treatise on the distribution of the fishes. In fact, his

publications are the only ones which deal in a comprehensive way with

the great mass of these fishes, including as they do almost two thousand

species.

In the problems of the origin and dispersal of Cichlid and Characinide,

Professor Higenmann (1906) has given interesting data. He has prepared

a hypothetical map which indicates that the Cichlid dispersed from east-

ern Guiana and the Characinide from the Amazon. This conclusion ap-

pears to be based on the fact that the most of the genera of Cichlide and

Characinidz are found in these regions. As I have already stated, how-

ever, this is no evidence for the point of origin and subsequent dispersal

of a family of animals. If it is, we might erroneously conclude from the

present distribution that the deer and tapir originated in the state of

Matto Grosso, Brazil, and the camel in the Andes, because more of the

species are found there; but in these latter cases, paleontology has shown

that the first point of origin was in the northern hemisphere, where the

species no longer exist. I hope to show in the following pages that the

point of origin of many species and genera of living Cichlidz has been in

the Amazon, but that the point of origin of the ancestral Cichlide was

not in South America.

In another paper, Professor Eigenmann draws the following conclu-

sions in his extended discussion of the distribution of the South American

fishes : :

1. The fishes of South America exhibit no close affinity with those of

North America.

2. The South American fishes, certainly the Characinide and Cichlid,

lend support to the Archhelenis theory.

3. The fishes of the coastwise streams of eastern Brazil differ more

widely from the Amazonian than do the Paraguayan.

4. The distribution of the fishes indicates that South America was

D2 \ ANNALS NEW YORK ACADEMY OF SCIENCES

‘divided into a northern and a southern part, 1. e., Archiplata and Arch-

-amazonia of von Ihering, and a connection appears to have existed be-

‘tween Guiana and Africa.

There may be little reason to question the data of Professor Higen-

‘mann, but there is, I believe, much reason to question his interpretations.

‘The first conclusion, we believe, is to be accepted for the living affinities

of the South American fishes, but when we are dealing with their points

of family origin, we are in most cases concerned with ancestral forms

“which have been dead for geological ages. In this case, the fossil record,

‘seanty though it be, shows affinities between the hving South American

cand the fossil North American fishes.?°@

Conclusion number two is also questionable, because:

‘1. There is no geological support for the Archhelenis theory. This is

all the more true for the late Cretaceous, when the Cichlide probably

originated.

2. The point of origin and dispersal of the Cichlids, as I propose to

show in the following pages, was not correctly determined.

In the matter of his third conclusion, it should be said in Professor

Eigenmann’s behalf that he did not then know that the Paraguay was not

connected with the Guaporé and that the Sao Francisco was connected

with the Tocantins. He also did not know which waterfalls were not

barriers for fishes and that both the coastwise streams and the Alto Rio

Parana have more than double the number of species which he assigned

to them.

In the same report, Professor Higenmann also states that, of the number

of species of fishes, 60 per cent of the Guianan, 40 per cent of the Sao

Franciscan, 53 per cent of the Paraguayan, 30 per cent of the coastwise

streams of eastern Brazil, 42 per cent of Trinidad and 6 per cent of Cen-

tral American are Amazonian. In this static comparison, he has intro-

‘duced a probable source of error due to the environments when he draws

conclusions from the above data. His lists include the fishes from the

entire basins of the coastwise streams, including Rio Sao Francisco, the

entire Amazon basin and only the central and upper Paraguay River,

which is only one of the affluents of the great La Plata basin. If we com-

pare the massive Amazon with the entire basin of the coastwise streams,

as 1t not necessary to compare the entire a Plata basin and not only one

part, 7. e., only one environmental complex? For example, if we should

compare the fishes from Rio Sapao of the Rio Sao Francisco with those of

the mighty Amazon and its affluents, 100 per cent would be Amazonian,

2a See H. F. OsBorn: “The Age of Mammals.” New York. 1910.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 53

and hence it would be more like the Amazonian than are the Paraguayan ;

but in this case we would compare only one environmental complex which

is duplicated in the enormous Amazon Valley. (In cases lke Central

America, the zodgeographer is concerned more with the ancestral dis-

tributed form than with the recent cenogenic modifications of them due

to the environment. )

In the same lists, we find 122 species for the lower Parana, only 64 of

which, or 52 per cent, are Paraguayan. If this could be true, then the

Paraguayan fishes are more like the Amazonian than the Paranean. This,

of course, is to a certain degree quite absurd, because small ocean steamers

can sail up the La Plata into the central course of either Rio Paraguay or

Rio Parana. In this case, according to the old view, fishes would, in the

first place, have to find their way overland from the Amazon Valley to the

Paraguay (a distance of about 200 miles separating the typical fauna of

the two basins), and then, for some unknown reason, remain in the Alto

Rio Paraguay and not venture to swim down the Paraguay into the

Parana.

Professor Eigenmann also states that the Alto Rio Parana had only 31

species, but I have often collected more than this in a single day. I col-

lected more than 100 species in the region of the Alto Rio Parana and its

affluents, and there is no reason to believe that the list was then exhausted.

The point brought out in the above brief review is that far too little is

known about the fishes or any other South American fauna to prove any

hypothesis by a numerical comparison of the species found in diverse

regions.

If, for example, we add Cichlasoma bimaculatum and Ovenicichla lepi-

dota to the Sao Franciscan fauna and Gymnotus carapo, etc., to the coast-

wise streams, and if we compare the entire river basin, there can be no

doubt that some of the faunal regions, and especially the cause of the

differences in their fauna as explained by Professor Higenmann, do not

agree with the actual facts.

The alleged support derived from the fishes for an Archiplata, Archi-

guiana and Archamazonia needs no discussion, because the geological evi-

dence shows that no post-Paleozoic seas have invaded the Plano Alto. In

the case of Patagonia, there have always been two possible connections

with the Plano Alto, one by the Cordova and the other by the Archean

rocks of the Andean region. Inasmuch as the invasions of the sea were

usually north and south, there is no evidence that southern South America

was completely isolated for a long period from the rest of South America

by an arm of the sea.?¢

°6 See PILsBry, 1911.

54 ANNALS NEW YORK ACADEMY OF SCIENCES

Use of Characters

The problem of defining the characters or the kind of characters

which distinguish species, genera, etc., has never been satisfactorily

solved. The writer will venture to consider only that part of it which

concerns the distribution of South American fishes.

In the main, there are two schools of ichthyologists, the American

and the English. The American school makes many divisions of the

families, genera and in some cases the species. ‘The English school has,

as a rule, been more conservative with taxonomic divisions and has

therefore fewer but larger groups. In support of the American school

may be given the results of the excellent experimental evidence obtained

by de Vries, Tower, Johannsen and others, which indicate that the sys-

tematic species is a complex one. In other words, this experimental

evidence tends to split up the species of the systematist into several ele-

mentary species. Much can be said in favor of this finer analysis of

species from the experimental standpoint, but little can be said in favor

of it from the standpoint of the systematist, because he does not know

whether his specimens are hybrids, whether they have a wide range of

fluctuating variation, whether they are mutations or whether the pecu-

liarities of the observed somatic differences are inherited or not.

Therefore, from the standpoint of geographical distribution, it ap-

pears that the English system, with its fewer divisions and divisions

based on more than single characters, is the better one, at least until we

have analyzed our species experimentally.

In reference to what characters are important from the standpoint of

the fish geography of South America, we are exceedingly fortunate, at

least in the case of Priscacara, a fossil cichlid described by Cope from

the Eocene of Green River, Wyoming.?’

From this interesting genus and from a comparative study of the

South American Cichlid, we are able to state with a high degree of

certainty that the ancestral Cichlide had the following characters:

Three anal spines; short gill rakers; more than one row of short conical

teeth in each jaw; pharyngeal teeth; ctenoid scales; serrated preoper-

culum; a continuous spiny and rayed dorsal with more than eight

spines; single naris or a tendency for narial coalescence; a rather short,

deep body, and a tendency to form a two-parted lateral line.2*

*7T have examined some of Cope’s types and believe that Woodward and Pellegrin are

correct in considering Priscacara a fossil cichlid.

28 Jt must be granted that it is difficult, if not impossible, to decide in the case of all

of the characters of fossil and living forms which characters are paleotelic and which

are cenotelic, but we can agree on at least a sufficient number to show that no living

cichlid fish could have given rise to those of both Africa and South America.

HASEMAN, GHOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 55.

These, then, were actually some of the characters of the primitive

Cichlide of the eastern and western hemispheres. |

These ancestral, primitive or phylogenetic characters may be desig-

nated paleotelic, a term which has already been used in a similar sense

by Gregory in his book on the orders of mammals.

In contrast to these paleotelic cichlid characters are the four to thir-

teen anal spines of Cichlasoma and other genera, the long gill rakers of

Chetobranchus and Chetobranchopsis, the lobe on the upper branch of

the first gill arch of Geophagus and Heterogramma, the long teeth of

Ptemia, the chisel or incisor teeth of Uraru, the long and more slender

body of Crenicichla, etc. These recent, adaptive or physiological char-

acters may be designated cenotelic. ‘There is no evidence that any of

these cenotelic characters have been distributed anywhere excepting in

South and Central America, because it is these and other characters

which distinguish the Cichlide of the western hemisphere from those of

the eastern.

I cannot overemphasize the importance of paleotelic and cenotelic

characters, because many zodlogists and paleontologists have not made

any distinction between these two types of characters in their tabulated

comparisons of various faunal regions. In the case of cichlid fishes,

cenotelic characters have evidently to do with the origin and dispersal of

variation, species, etc., while paleotelic characters deal with the ancestral

fauna which gave rise to these genera. The paleotelic characters have

to do with the ancient distribution, hence theories like Archhelenis;

while the cenotelic have to do with the present distribution of a given

genus. ‘The cenotelic characters are usually modified by the action of

the environment on the ancestral forms of a given genus, while the paleo-

telic characters in part have extended down through all of the genera of

the cichlid family. Therefore, from the standpoint of the origin and

lines of dispersal of the Cichlids, a few paleotelic characters will out-

weigh a bookful of cenotelic ones.

What shall we learn, then, by a careful compilation of all of the Cich-

lide of South America and Africa and by comparing all of those found

m one river basin with those found in another? Would this show that

a connection had existed between certain points of Africa and South

America? Or would it merely be a compilation of cenotelic characters

formed by the action of the environments of the different localities on

the ancestral Cichlide which possessed paleotelic and not the recent

cenotelic or secondary characters ? ‘

From the standpoint of the origin of the ancestral Cichlids, then, the

living species or the specific characters alone give us no clue, because

56 ANNALS NEW YORK ACADEMY OF SCIENCES

they are based on cenotelic characters. The generic characters are often

no better, because they, too, are usually cenotelic.

Take, for example, the genus Cichlasoma, which is primarily distin-

guished by the presence of four to thirteen anal spines from the genus

Afquidens, which has only three anal spines. For all we know, a muta-

tion with four anal spines could have easily appeared from the three anal

forms, or vice versa. In fact, that is exactly what seems to be the case

in Rio Sao Francisco, where I found only two specimens with three anal

splines among thousands of the four-anal-spine form, Cichlasoma bimac-

ulatum. In southern Brazil, I found just the opposite, namely, only

three four-anal-spine forms of the same size, color, etc., as thousands of

three-anal-spine forms which are known as Avquidens portalegrensis.

Hence all such closely drawn genera must be used with great care in the

determination of the point of family origin and dispersal, because they

are based on such characters that a mutation or individual or local varia--

tion might readily establish a new genus and species in a very short time.

The importance of having a clear idea of the exact status of the ich-

thyological taxonomy, and especially the value of generic and specific

character and which are paleotelic and which are cenotelic, are of prime

importance in the explanation of the distribution of the fishes. For ex-

ample, the writer described Geophagus brasiliensis iporangensis as a new

variety from the headwaters of Rio Ribeira de Iguape. This variety may

be a distinct species, and it may be only a somatic change due to the en-

vironment. ‘The adult forms certainly look and measure like a distinct

species, but I observed that the young individuals, one and two inches

long, could not be told from the lowland young of Geophagus brasiliensis.

It is highly probable that this new variety would produce typical Geo-

phagus brasiliensis, if it were removed to the lower course of the same

river. ‘The same may be true of Crenicichla iquassuensis, also described

by me from the Rio Iguasst. This species is closely related to C. lacus-

iris and would perhaps produce that species, if subjected to the same ex-

ternal conditions. These two examples illustrate the present status of

South American ichthyology. One can, I believe, readily separate out a

couple of hundred forms of the two thousand catalogued for South

America which are probably not species, but merely “ontogenetic species,”

showing, for example, somatic changes of a color spot, a few more or less

scales and spines or some other trivial physiological difference. Many

of these species, in fact, are based on single characters, such as teeth,

which are variable structures. The same have been shown experimentally

with birds, beetles, butterflies, etc., ving in different temperatures, etc.,

and until the fishes are better and experimentally known, their present

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 57

static distribution offers only hypothetical evidence of the shakiest kind

for any theory.

Ichthyological Faunal Regions

In the first part of this thesis, the writer briefly outlined the principal

environmental complexes in which the sum total of the natural condi-

tions was about equal in each complex. In the following pages, I pro-

pose to analyze the changes which have been wrought by the environments

on the distribution of the fishes: but before making the analysis, it is

first necessary to consider the range of the fishes.

The Siluride are found everywhere in South America. The Cichlidee

and Characinide are found everywhere north of Patagonia and east of ©

the Andes.. A few Characinide and Cichlid are found on the western

slope of the Andes of Peru and Central America, and a few are also

found in the West Indies. Two species of Characinidz are also found in

northern Patagonia.

The few species found in the West Indies diminish in number of

genera the farther the islands are from the mainland ‘There is thus a

suggestion that the Cichlide and Characinide came directly from South

America by way of the sea. I have no doubt that this is true, since by

actual experiment I have determined that certain of these genera will

live for some time in sea water.

The Characinide have never been completely revised, and consequently

the genera cannot be of equal value. Notwithstanding this lack of re-

vision, the following summary of the genera is instructive. Of the 129

genera of Characinide which have been described, 94 are found in the

Amazon Valley, 50 are widely distributed, 60 are found in the Paraguay,

64 in the Guianas, 58 in the Orinoco, 54 in the La Plata, 41 in the Sao

Francisco and 37 in the coastwise streams of southeastern Brazil. The

_ Paraguay harbors about 60 genera of Characinide, 58 of which are found

in the Amazon. The species belonging to these genera are not usually as

similar as are the genera, for example, of the 118 species of Characinidee

reported by Higenmann from the Paraguay, only 63 are found in the

Amazon, while 45 of his 47 Paraguayan genera are found in the Amazon.

Do the above data indicate a direct connection between the Paraguay and

the Amazon? No, and for the following reasons:

1. There is at the present day no connection and no indication of an

ancient connection, at least since the present ichthyological fauna has

developed, at such an altitude as to be favorable for lowland forms to

cross from one basin into the other.

2. No connection is known to exist between the rivers of Guiana and

58 ANNALS NEW YORK ACADEMY OF SCIENCES

the Amazon, and yet there is as great, if not greater, identity of fishes

there than in the case of the Paraguay.

3. There are connections between the Orinoco and the Amazon and

the Sao Francisco, and yet there is less similarity between their faunas

than between these of the Paraguay, Guiana and the Amazon.

The key to the explanation of the distribution of the characinids lies in

the 50 genera which are not only widely distributed, more or less cos-

mopolitan, but the more generalized members of their families. The

same is also strongly indicated by the fact that when these widely dis-

tributed generalized genera arrived in the different environmental com-

plexes, there resulted less conformity between the species than between

the genera and a greater similarity between the species which lived in

similar environments, even though they are not connected, than between

the species in dissimilar environments which are connected.

I have already shown that the Plano Alto separates the La Plata basin

from the Amazon. This being the case, we have to look for another ex-

planation of the distribution of the fishes other than river connections.

The most natural way to seek the explanation is first to consider the

fishes which live on the Plano Alto.

I have found the following genera on the Brazilian highlands (a max-

imum list) :

Siluride:

Callichthys, Pydidiwm, Rhamdia, Pimelodus, Pimeladella, Plecostomus,

Doras, Trachycorystes, Auchenipterus, Heleogenes, Loricaria, Hoplos-

ternum and Corydoras.

Characinide :

Erythrinus, Hoplias, Holerythrinus, Characidiwn, Oreatochanes, Pecilu-

richthys, Acestorhynchus, Curimatus, Moenkhausia, Astynax, Tetrago-

nopterus, Phyrrhulina, Pecilocharaxz, Serrasalmo and Chalcinus.

Cichlide:

Geophagus, Crenicichla, Aiquidens, Cichlasoma and in two places Hereto-

gramma.

Rivulus, Symbranchus marmoratus, Hypopomus brevirostris, Gymnotus

carapo, Higenmannia virescens and Sternopygus macrurus.

All of the above genera are not only widely distributed but also the

most generalized types of their sub-families. They are represented in

the highlands proper by only a few species. For example, I found only

twenty-five species in the highlands of Parana above the big Iguassu

Falls. The following is the list which I collected in the highlands of

northern Goyaz:

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 59

Hoplias malabricus, Bloch.

Acestorhynchus falcatus, Bloch.

Hoplerythrinus uniteniatus, Spix.

Characidium fasciatum, Reinhardt.

Curimatus elegans, Steindachner.

Creatochanes sp.?

Moenkhausia oligolepsis (?) Gunther.

Astaynax bimaculatus, Linnzus.

Crenicichla lepidota, Heckel.

Cichlasoma bimaculatuwm, Linnzeus.

A similar paucity of species has been noted by Eigenmann in the high-

lands of Guiana. He has kindly given me the following list which he

collected above the Kaieteur Falls:

Rhamdia quelens Pecilurichthys bimaculatus

Heleogenes marmoratus Astyanax mutator

“Pydidium gwianense Hoplias malabricus

Callichthys callichthys Hoplerythrinus uniteniatus

Lithogenes villosus Erythrinus erythrinus

Corymbophanes andersoni Gymnotus carapo

Phyrruhlina filamentosa Hypopomus brevirostris

Pecilocharazx bovalli Rivulus holmie

Moenkhausia oligolepis Ajquidens potarensis

Moenkhausia brown Heterogramma ortmanni

Creatochanes affinis Crenicichla alta

It is at once evident from the above that there have never been found

more than twenty-five species in any one locality on the Plano Alto.

Compiling the possible lists from various localities, however, we might

obtain as many as fifty species as inhabitants of the Plano Alto. If we

even doubled this probable list, there would still be about fifty species,

many belonging to lowland genera common to the Paraguay and Amazon

valleys, to account for in some other way than by a possible overland

passage. Hence other factors than mere land-bridges, river connections,

etc., are involved in the present distribution of South American fishes.

It has already been stated that the highland genera, usually small in

size and widely distributed, are the generalized types which have pro-

duced the bulk of the ichthyological fauna. In fact, the highland genera

which I have enumerated include 33 per cent of the 1917 species which

have been reported from South and Central America by Higenmann.

The highland genera of Cichlid include 150 of the 187 species of Cich-

lide reported by Eigenmann from South and Central America.

The above statistics are sufficient to show that most of the purely fresh-

water fishes are directly related to the highland genera which continue to

60 ANNALS NEW YORK ACADEMY OF SCIENCES

enter practically all of the rivers north of Patagonia and east of the

Andes. None of these highland genera need a direct connection between

the river basins, because they are found not only above high waterfalls -

which have been barriers, but also in all of the river basins, some of which

are distinctly separated. In this connection, I may note that I saw young

Hoplias swimming during a heavy rain in a trail over the highlands of

northern Goyaz fully two miles away from the nearest rill.

I have already stated that the highland genera directly account for the

distribution of at least 33 per cent of the entire ichthyological fauna

without the necessity of direct connections between the different river

basins. Neither river connections nor the existing highland genera,

however, will account directly for all of the species in common between

any of the rivers; nor will they explain the more difficult question why

so many of the common species have remained identical in river basins

which may or may not be connected. In order to answer this most diffi-

cult question, I have chosen the cichlid fishes, because they are the best

known of any large family of South American animals.

In brief, the question is, Why are there more species of Cichlid in

common in Guiana, the Paraguay and the Amazon than in the Parana,

Uruguay, the coastwise streams of southeastern Brazil aad the Amazon?

The following list gives the genera of Cichlide and their distribution.

Those marked with an asterisk (*) have few species and are more closely

drawn than the others. The word “general” means everywhere north of

Patagonia and east of the Andes.

* Chetobranchus, Amazon to North,

* Chetobranchopsis, Amazon and Paraguay,

* Cichla, Amazon to north, Orinoco and Guiana,

* Uraru, Guiana and Amazon,

* Herotilapia, Lake Managua,

* Neetroplus, eastern slopes of Mexico and Central America,

* Acaropsis, Amazon, Guiana and Orinoco, :

* Petenia, Lake Peten,

* Tomocichla, Costa Rico,

Herichthys, Texas to Guatemala,

* Astronotus, Paraguay, Amazon and Orinoco,

* Vannacara, Essequibo,

Ajquidens, general and western Ecuador,

Thorichthys, eastern slopes of Mexico and Central America,

Cichlasoma, general and both slopes of Central America,

Crenicara, Amazon and Guiana,

Crenicichla, general,

* Retroculus, Amazon,

Heterogramma, Paraguay, Amazon and Guiana,

Geophagus, general,

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 61

* Symphysodon, Amazon,

* Pterophyllum, Amazon, Guiana and Orinoco,

* Biotecus, Saraca in Amazon, ;

* Paranecetroplus, Rio Sarabia, Mexico.

According to the static viewpoint of animal geography, we shouid con-

clude from the above data that the Cichlide originated or dispersed from

either the central or the northern part of the Amazon Valley, because

sixteen of the twenty-three genera are found there. Five other genera

are found only in Central America. All of the seven genera which are ©

found south and east of the Amazon basin are found also in Rio Ama-

zonas. Statically, also, we could interpret the entire absence of Cichlidze

from Patagonia as meaning that this group had a more northern origin;

but inasmuch as the genus A’quidens®® possesses more of the paleotelic

characters of the ancestral Cichlid, it is evident from its distribution

that it may have originated in many places not embraced by the mighty

Amazon. Also the fact that at least eight of the sixteen Amazoniam

genera are highly specialized, 7. e.. cenotelic, may be taken as evidence

that these forms have evolved in this region from less specialized forms

whose center of origin was not necessarily in the Amazon. In fact, the

Amazon has only two genera which are not found elsewhere in South

America, and both of these genera are closely drawn and contain only one

species each.. Therefore the above list of genera and all that is known

about them offer no conclusive evidence that the majority of the living

American Cichlid originated in the Amazon. They may equally well

have origimated, as far as the above evidence shows, in either Guiana, the

Ormoco or anywhere on the old Plano Alto. Before attempting to deter-

mine the poimt of origi of the Cichlidx, I will first put some of the

typieal genera im their environmental complexes in order to explain their

present distribution: iA

Rro Uruguay anp Rio GRANDE DO Sut, INCLUDING PART OF THE LOWER

La PLAtTs

This complex. is characterized by medium to low altitudes, sub-tropical

to temperate climates, campos, slow flowing water and little or no forests.

This region harbors the following species of Cichlide:

23 Pellegrin (1904) considers this the most primitive genus, but Crenicara also has:

many paleotelic characters.

6—NY

62 ANNALS NEW YORK ACADEMY OF SCIENCES

( balzanii,

Geophagus brasiliensis / brachyurus,

gymnogeyns,

brachyurus,

balzanii,

Aiquidens portalegrensis — Cichlasoma bimaculatum, haying only one

anal spine more,

Cichlasoma facetum,

Crenicichla lepidota, southern form of C. saxatilis,

lacustris,

vittata, southern form of C. macrophthalmus.

These species are found on both sides of the divide in the State of Rio

Grande do Sul. The region is characterized by Geophagus gymnogeyns

and brachyurus.

ALTO PARANA AND COASTWISE STREAMS OF HASTERN BRAZIL

This complex is characterized by higher altitudes, numerous water-

falls, rapid water, sub-tropical temperature and forests. It harbors the

following species of the Cichlid:

Geophagus brasiliensis with two varieties,

Bis 3 CO. jaguarensis

Crenicichla lacustris i 0 eee

& lensis,

vittata (C. dorsocellata, southern form of C. macrophthalmus),

lepidota, southern form of C. sazxatilis,

Cichlasoma facetwin (C. autochthon and C. oblongwm being synonymous),

Cichlasoma bimaculatum, northern form of A/quidens portalegrensis.

These six species and their varieties are found on both sides of the

divides between the headwaters of the Alto Parana and the coastwise

rivers of eastern Brazil. This region is characterized by two varieties of

Geophagus brasiliensis, two varieties of Crenicichla lacustris and one

variety of Crenicichila vittata. Whether these varieties will breed true or

whether they exhibit only somatic changes which are not necessarily

inherited is not known, but they show, at any rate, changes due to the

peculiar environments in which they live. For example, the young of

Geophagus brasiliensis var. iporangensis could not be distinguished from

the young of the common form, but the adults were strikingly different.

The variety lived in the rushing headwaters of Rio Ribeira and the com-

mon form, G. brasiliensis, lives more in the lowland sections of the rivers,

lagoons and swamps. I observed a similar change from lowland to high-

Jand forms near Santos. I also observed that Geophagus brasiliensis

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 63

occasionally is taken from salty water. I put this species directly out of

fresh into a bucket full of sea water. It was able to live several hours

under these conditions.

Sao FRANCISCO AND THE Secca (Dry) REGion or NorTHEASTERN

BRAZIL

This region is characterized by desert-like flora, no forests, scanty rain-

fall and occasional long dry seasons during which many rivers become

dry. The altitude is medium (246 m. at Jatoba above the Paulo Affonso

Falls).

This region harbors the following species of the cichlid fishes :

Cichlosoma bimaculatum,®

Crenicichla lepidota,

Geophagus brasiliensis.

It is interesting to note that the above three genera are also the only

three which are found in Rio Grande do Sul, where, in place of having

the same three primitive species, six new more cenotelic species have

evolved from the above three more generalized widely distributed forms.

The only explanation is that the environment of the muddy semi-arid

Rio Sao Francisco has not been conducive to either the production-or the

maintenance of new species, because the original species were identical

for both of these environmental complexes, i. e., they came from the same

ancestral highland stock.

THE PARAGUAY AS PaRT OF THE AMAZONIAN COMPLEX

The great swamps, called pantanals, of the Paraguay are in all respects

the exact southern counterpart of part of the Guaporé and the central

portion of the Amazon basin. Their similarity must be great, because

they all lie in the confines of the central portion of the remains of the

Plano Alto. This is not true of either the Parana, Uruguay or Sao

Francisco rivers, as well as the Andean affluents of the Amazon. The

similarity is further very striking in altitude, while the Parana is three

or four times higher. Inasmuch, however, as the Amazon Valley is so

large, it duplicates several times the natural conditions found in the

Paraguay as well as in several other rivers. This duplication includes

temperature, altitude, food, volume of water, swamps, nature of currents,

humidity, rainfall, nature of sediment and muddy and clear water.

°°T collected two specimens with three anal spines which were exactly like Hgwidens

portalegrensis.

64 ANNALS’ NEW YORK ACADEMY OF SCIENCES

Given then this duplication of environmental complexes and given also

the same common generalized widely distributed genera of Cichlid, may

we not also expect some similar changes in the common germplasm?

The Paraguay harbors the following species of cichlid fishes:

Chetobranchopsis australis, southern form of C. orbicularis,

Astronotus ocellata,

Alquidens paraguayensis, southern form of A. teteramerus,

portalegrensis, southern form of Cichlasoma bimaculatum,

dorsigera,

Cichlasoma festivum,

Orenicichla simoni, perhaps synonymous with C. reticulata,

semifasciata, perhaps Synonymous with C. cyanonotus,

lepidota, southern form of C. saxatilis,

vittata, southern form of C. macrophthalmus,

Heterogramma teniatum,

trifasciatwn,

borelli, giving off H. ritense,

corumbe,

Geophagus balzanii,

jurupari.

Of the sixteen species of Cichlid, only five are found in Rio Uruguay

and Rio Grande do Sul, only three are found in Rio Sao Francisco and ~

only three are found in the Alto Rio Parana and the coastwise streams of

southeastern Brazil. These three are widely distributed species. All

of these sixteen species excepting five are also found in the Amazon, and

of these five, two are Heterogramma, which are connected by intermediate

stages in such a way that Heterogramma teniatum can easily give rise to

all the species of this genus. Two of the other species which are not

found in the Amazon are Crenicichla semifasciata and simon, but these

are still questionable species, and they also may even exist in the Amazon.

In order to explain this identity of the Cichlide, I might even grant that

all of these species have in some unknown way interchanged between the

Agpazon basin and the Paraguay. If, however, I should grant so much

as that, I should then find even greater trouble in explaining not only

why the rest of the Cichlide of the Guaporé were not able to get in the

Paraguay, but also why the Paraguayan species have remained identical

with those of the Amazon and why the Paraguayan species have not in-

vaded the Rio Parana and Rio Uruguay, all of which have navigable

channels in their lower courses. In other words, the La Plata basim has a

triple cichlid fauna, and these correspond exactly with the natural en-

vironments of the Alto Parana and the coastwise streams of eastern

Brazil, Kio Uruguay and Rio Grande do Sul, and Rio Paraguay The

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 65

changes wrought by these environments on the more generalized high-

land genera is adequate, I believe, to account for the present distribution

of the Cichlide, but this does not explain their point of origin.

Before taking up this latter subject, it is necessary to produce further

evidence showing that it is the action of the environmental complexes on

widely distributed genera which has produced the present distribution of

the South American fishes and not direct river connections or inter-

mingling of species and isolation.

Rio AMAzONAS REGION

The following is a list of the Cichlid of Rio Amazonas :*+

Chetobranchus flavescens, Guaporé,

semifasciata,

- Chetobranchopsis orbicularis,

Cichla temensis, Orinoco,

ocellaris, Orinoco, Guiana, Guaporé,

Uraru amphiacanthoides,

Acaropsis nassa, Guaporé,

Astronotus ocellaris, Guiana, Orinoco,

orbiculatus,

Ajiquidens teteramerus, Hssequibo, Guaporé,

vittata, Colombia, Guiana,

paraguayensis, Guaporé,

subocularis, Guiana,

portalegrensis, Guaporé,

dorsigera, Guaporé,

duopunctata,

zamorensis,

guaporensis, Guaporé,

awani, Guaporé,

Cichlasoma bimaculatum,

festivum, Guaporé,

severum, Guiana, Guaporé,

psittacum, Orinoco,

spectabile, .

coryphenoides,

Crenicara altispinosa, Mamoré, -

maculata,

punctulata, Guiana,

i This list does not exactly agree with my report from the Carnegie Museum, which

was slightly changed by Professor Eigenmann. Perhaps somé of the omitted species

should be added, but I am inclined to believe more should be dropped, even @ few de-

scribed by the writer. In the main, however, this is the most accurate list at hand and

is sufficient for its present purpose, including as it does all the genera.

66 ANNALS NEW YORK ACADEMY OF SCIENCES

Crenicichla reticulata, Guiana,

cyanonotus,

lepidota, Guaporé,

saxatilis, to the north,

lucius, Guiana,

macropthalmus, Guaporé,

acutirostris,

lenticulata, Guiana,

strigata,

cincta,

johanna, Venezuela, Guiana, Guaporé,

lugubris, Venezuela, Guiana, Guaporé,

santeremensis,

Retroculus ladifer,

Heterogramma teniatum and a variety, Guaporé,

agassizi, Guaporé,

trifasciatum and a variety, Guaporé,

corumbe, Guapore,

Geophagus surinamensis, Guaporé,

cupido, Essequibo,

jurupari, Guapore,

acuticeps.

Biotecus opercularis,

Symphysodon discus,

Pterophyllum scalare.

It may at first sight appear strange that the Amazon harbors so many

species and genera of Cichlid, but it is exactly what one would expect

because of its vast size and tropical location. Of these fifty-three species,

at least twenty-two are found in Rio Guaporé, and of these twenty-two,

twelve are not found in the Paraguay. This fact alone is sufficient to

disprove any wholesale exchange of fishes between the Paraguay and the

Guaporé. From the standpoint of phylogeny, I am able to throw a little

light on the above distribution of the Cichlide. These conclusions were

derived from both the field and laboratory, and inasmuch as the purely

systematic data have already been published by Pellegrin, Regan and

more recently by myself, I will not repeat them.

1. I consider Geophagus brasiliensis as the most primitive of the living

members of this genus. It is interesting to note that this species is not

found in the Amazon and that its nearest northern ally is Geophagus

steindackneri, which was originally described by Steindachner as Geopha- -

gus brasiliensis from Rio Magdalena, which, like the coastwise streams

of southeastern Brazil, flows out of Archean mountains and possesses

therefore remarkably similar environments.

2. Aiquidens teteramerus appears to be the most primitive of its genus.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 67

It changes in the Guaporé River into A. paraguayensis, which is also

closely related to A. vittata. All of the species of this genus form a

natural group about A. teteramerus, excepting A. portalegrensis, and I

consider it as giving rise (or vice versa) to Cichlasoma bimaculatum.

3. Heterogramma tematum can easily give rise to all of the species of

this genus. In fact, I have reasons to doubt the reality of all of these

species, because they may be nothing more than fluctuating variations,

principally in color, or somatic changes which may or may not be in-

herited. At any rate, there is an almost complete intergradation of all of

the species of this genus. Hence experimental work is needed before this

genus can be properly classified.

4. There can be no doubt that Cichlasoma bimaculatum is the most

primitive of its genus, because it is not well defined from Mquidens

portalegrensis, which is the most primitive living cichlid genus.

5. I consider Crenicichla saxatilis as the most primitive of its genus.

It is represented in the south by QO. lepidota. C. vittata is the southern

form of C. macropthalmus, and they are connected by varieties through

C. lucwus to saratilis. C. johanna and the other elongate Amazonian

species of this genus also can be linked to C. savatilis. |

I will not venture to discuss the relationship of the other genera and

species, because the results would be only an opinion with little or no

support. The above brief consideration, however, is extremely ‘useful,

because Cichiasoma bimaculatum, Aiquidens teteramerus, Geophagus

brasiliensis and Crenicichla saxatilis are the generalized types which not

only are widely distributed but also have been the origin of the bulk of

the Cichlids found in the various environmental complexes. The genus

Cichlasoma alone, according to Eigenmann, contains eighty-four of the

one hundred eighty-seven known species of American Cichlid. These

four genera actually embrace at least 80 per cent of the species of the

American Cichlide, and several other genera can easily be derived from

them. i

These four genera are found from one end of the Plano Alto to the

other, and consequently from their present distribution we can explain

the origin and distribution of their derivatives, but this has nothing to

do with the origin of the cichlid family.

To sum up briefly, then, the distribution of the Cichlide, we may say

that three highland genera are found in Rio Sao Francisco and have not

evolved any new species. The same three genera have produced nine

species in the Rio Grande do Sul, 7. ¢., the three old plus six new species,.

and six species in the Alto Rio Parana and the coastwise streams of east-

ern Brazil. Sixty per cent of the Paraguayan Cichlide are also included

68 ANNALS NEW YORK ACADEMY OF SCIENCES

‘by the same three genera and more than 50 per cent of the Cichlid of

the Guaporé are not found in the Paraguay.

- The diagram forming Plate XVI indicates the evolution and distribu-_

ction of the-cichlid fishes of the Amazon Valley and the rivers south of it.

dt shows that river connections or interchanging of fauna and barriers

or isolation are not the important factors of geographical distribution,

but that the organic complex of the ancestral stock (three highland gen-

era—Geophagus, Crenicichla and Alquidens-Cichlasoma) and the compo-

sition of the environmental complexes in which they came to live, pro-

duced by the rivers sinking into the Plano Alto, are the important factors.

The figure also shows that similar and identical evolution of the common

ancestral stock has taken place in similar environments and dissimilar

evolution in dissimilar environments regardless of whether the environ-

ments are or are not connected. (See Plate X V.**)

The phylogeny of the extra Amazonian species, 7. e., more than thirty-

three (fifty-three existing in it), is not yet clear, but they will eventually

be deduced from the highland stock, because I have shown that the

Amazon Valley as we now know it has existed a comparatively short

time.

The facts as shown on the diagram are almost exactly the opposite to

what one would expect, if land and water connections or isolation were

the important factors of living animal distribution. The numbers at

the end of the arrows show the number of new species which have un-

questionably descended from the old highland stock when it entered the

rivers which were gradually eroded in the Plano Alto. The diagram

also shows the basins connected or not and the identity of fauna in dis-

connected regions, such as Alto Parana and coastal streams. I have no

first hand knowledge of the Rio Orinoco; hence I do not discuss it.

Additional proof of similar evolution in similar environmental com-

plexes, even if they are not connected, is offered by the larger species of

South American fishes. For the sake of clearness, I have divided a

typical abbreviated list of large species of fishes into the following

classes :

: 1. Large species of fishes found in the upper Guaporé and Amazon,

and neither the genera nor the species found in the Paraguay-La Plata

basin. A few examples of such fishes are Cichla ocellaris, Phracto-

cephalus hemiliopterus, Brachyplatystoma reticulatum and Electrophorus

electricus. 'To these and many other fishes may be added the large croco-

82 Whether we should call these isolated species identical or by the same name may be

a debatable question. Not any two individuals are identical, but these species are not at

present distinguishable.

ANNALS N. Y. ACAD. SCI. VOLUME XXII, PLatTE XVI

OUTLINE MAP OF PART OF SOUTH AMERICA

Showing the centers of evolution and the distribution of the cichlid fishes of the Amazon

Valley and the rivers south of .it

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 69

dilian Caiman niger, the large Amazonian turtles Podocenemis expansa

and P. tracaxa and the red porpoise Inia geoffroyensis. All of these

forms are found as far up Rio Guaporé as Bastos, Rio Alegre. If any

direct swamp connections existed, I should certainly expect to find these

animals in the Paraguayan pantanals. ‘The smaller species of caiman,

Caiman sclerops, is found.all over South America excepting Patagonia

and west of the Andes, but it has been seen six miles away from water,

and hence an overland trip is not impossible for it.

2. Large species of fishes found in both the Guaporé-Amazon and the

Paraguay-La Plata, but never near their headwaters, 7. e., at least fifty

miles apart in a straight line between the two basins. A few typical

examples of such fishes are Sorubim lima, Hemisorubim platyrhynchus,

Sciades pictus, Mylossoma aureus, Charax gibbosus, the giant Paulacea

jahu of La Plata and Paulacea lutkent of Amazon (I consider the last

two species synonymous). . These and at least fifty other species which

are found both in the Amazon and the Paraguay have not interchanged

as such between these basins, for the following reasons:

1. No connection has existed.

2. They are not found in the headwaters, 7. ¢., above the waterfalls

and at such high altitudes as exist between these rivers.

3. The distance between the headwaters is so great that an accidental

distribution is not possible.

4. Hven if a connection had existed, it would not explain why the spe-

cies have remained the same and why Rio Uruguay and Parana, belong-

ing to the same river basin as Rio Paraguay, do not possess all of these

species but, on the contrary, harbor many species not found in Rio Para-

guay. Why, also, did not other species interchange, if a connection has

existed ?

5. The Sao Francisco River is connected with the Amazon Valley, yet

it does not have nearly as many Amazonian species as does the Paraguay.

In fact, its common or Amazonian species are cosmopolitan forms. Rio

Sao Francisco has an unfavorable environment (dry, hot, high, muddy,

fewer swamps, etc.) and therefore has fewer species than the large Para-

guay, with its favorable cichlid environments which have produced more

cenotelic changes in the ancestral stock.

Tn view of all this, it appears that the only answer which can be given

to the question why the Paraguay has at least 53 per cent of Amazonian

fishes is .,,. las ais

1. About 50 per cent of the similarity is due to the cosmopolitan forms,

1.-€., to-overland distribution of the small generalized highland genera

which are widely distributed.

70 ANNALS NEW YORK ACADEMY OF SCIENCES

2. When these highland forms arrived in the same kind of environ-

ments, they often underwent identical evolution with that which was

taking place somewhere in the massive Amazon.

3. The remainder of the similarity is due to marine immigrants.

The first part of this answer needs no further comment, but the second

may appear to be absurd, at least to those who are not familiar either

with the South American fishes or with the environmental complexes in

which these fishes live. The view that the common highland genera of

fishes have often undergone identical evolution in similar environments,

even if these environments are well separated, is of the same general

nature as those given in the following publications.

Bateson has shown in the case of Cardiwm edule of the Aral Sea that,

as the sea dried up, isolated basins were formed in which the salinity

was greatly increased, and under these conditions the cockles so separated

show similar variations under similar conditions. These shells of the

cockles in the higher to the lower terraces showed a progressive change

in regard to the following features:

(a) Shells became much thinner.

(b) Shells became highly colored.

(d) Shells became smaller in size.

(¢) The grooves between the ribs on the outside appeared on the in-

side of the shells as ridges with rectangular faces.

(f) A great increase in length in proportion to the breadth of the

shells.

Are not the changes observed by Bateson as profound as required to make

Cichlasoma bimaculatum out of Mquidens portalegrensis. or vice versa?

In this case, the loss or gain of one spine makes a different genus and a

species. ‘Thus it is with many other genera and species of fishes whose

distribution must be explained by identical evolution in similar environ-

ments.*8

MacDougal has obtained nearly all of the mutations observed by

de Vries in Holland from @nothera lamarckiana obtained in France,

England and Holland and planted in New York. He also obtained one

mutant, O. albida, from O. lamarckiana Nantucket City. Even if O.

lamarckiana is a hybrid, it makes no difference, for at least part of the

common highland stock may also be hybrid.

Tower has shown, both in nature and by experiments with humidity

and temperature on Leptinotarsa, that he could produce changes both in

83 Some of the small differences may be purely somatic, which are not necessarily

inherited. Wxperimental evidence is necessary to settle this point.

_. HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA “71

the germplasm and somaplasm or in each separately. For example, he

was able to increase the temperature in the laboratory at Chicago and

obtain forms of the potato beetle which lived in Mexico. Tower’s work

shows almost beyond a doubt that when the same species of potato beetle

lived in different environmental complexes, similar variations were pro-

duced in similar environments. This is all that is required to explain

the similarity of certain genera and species of fishes which are found in

the Paraguay and Amazon, but not in the Parana, Uruguay or the coast-

wise streams of eastern Brazil. Furthermore, the requirement is not

ereat, because many of the genera and species of fishes are based on one

more or less spine, three to ten more or less scales in the lateral line, color

and position of spots and other trivial characters which are subject to a

wide range of so-called fluctuating variation. In fact, some of these

variations may even exist occasionally in the generalized widely distrib-

uted highland species from which the bulk of the ichthyological fauna

of the various river basins has evolved.

Any one less familiar than the writer with the region in question would

not venture to state that identical evolution has taken place on such a

large scale in similar environments. As I have already shown, however,

no connection exists between the Paraguay and the Amazon. Accidental

overland and marine distribution is more absurd than identical evolution

from common highland stock, and even if the species got across, we

should still have to admit that they have remained almost the same in

the case of the Paraguay and the Amazon and have not in the case of

the Parana and Uruguay rivers, which belong to the same basin as the

Paraguay. To admit the latter is equivalent to admitting either that

identical evolution has taken place in the case of many genera and species

of fishes, or else to believing in the fixity of species in one locality and

not in another. Furthermore, the Sao Francisco has a connection with

the Amazon, and yet its cichlid fauna is composed of the three common -

highland genera and species only. If a connection is needed to explain

the similarity of the Paraguayan fishes with the Amazon, I desire to ask,

“Why has the Rio Sao Francisco only three species of cichlids? Why

is there a triple cichlid faunal region in the La Plata basin? Why have

the Paraguayan species remained identical with the Amazonian? Why

did not more of the Cichlid of Rio Guaporé enter the Paraguay? My

answer to these questions is that similar environments have produced

some similar changes in the same germplasm, 1. e., the highland genera

which are widely distributed,** and dissimilar environments have pro-

% This does not at all imply the inheritance of acquired characters, for it can also

easily be a direct effect on the eggs, sperms or germplasm.

72 ANNALS NEW YORK ACADEMY OF SCIENCES

duced some dissimilar changes. Many of these changes are of such a

nature that the species are adapted to live only in certain kinds of en-

vironments. The cichlid fishes usually live in swamps, lagoons or lakes,

and seldom in rapidly flowing water. In the open channels, the chara- -

cins would eat them. The Paraguay and the Amazon have many swamps

and many cichlid fishes. Alto Rio Parana has few swamps and few

eichhids.

The marine immigrants which have entered the rivers and become

permanent dwellers of the same have also increased the percentage of

similarity between certain rivers more than between others. Typical

examples of such fishes are as follows:

1. The fresh-water skates, Potamotrygon, two species of which are

found both in the Paraguay and the Amazon. We have no evidence that

these two species separately left the ocean and became dwellers of these

rivers. Other species of the same genus are found only in the Amazon,

in the La Plata, the Guiana and the Orinoco. Is it not possible that some

of these species are the results of changes invoked in the marine ancestor

by their new environments ?

2. Two species of Peeciliide, Rivulus and Girardinus, could easily have

gone along the coast from the La Plata to the Amazon. They may also

have gone overland.

3. Stoleophorus olidus.

4. Two species of Scieenide belonging to the genus Pachyurus.

5. Several catfishes, and even a few of the characinids and cichlids,

might have migrated along the coast, but it is out of the question to as-

sume that all of the identical forms in Rio Paraguay and Rio Amazonas

did so because they are not found in coastwise streams, Rio Uruguay and

Rio Parana.

There is no doubt that the marine immigrants have played an im-

portant part in the production of a greater similarity between the fresh-

water fishes of certain river basins than others. I found only two marine

species of fish in Rio Colorado of Patagonia and at least one hundred are

found in the lower Amazon. Even the sawfish (Pristis) is sometimes

killed as far up the Amazon as Santerem (476 miles).

The volume of water always bears a relation to the number and size of

marine immigrants, and this is especially true when there are many

islands, many channels, plenty of food, tidal effects and much brackish

water. :

It is evident, then, that the small rapid coastwise streams of eastern

Brazil, Rio Magdalena and Patagonia should have fewer species than

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 73

either the Amazon or the La Plata, regardless of any hypothetical con-

nection with the eastern hemisphere, for the following reasons:

1. Smaller volume of water, relative higher altitude, excepting parts

of Patagonia and strong currents.

2. The original stock, part of which was marine in origin, did not

develop into so many species and genera in the restricted environments

as it did in the more extensive environments, because the factors active

in the evolution and preservation of life were neither as favorable nor as

numerous in the restricted environments.

3. Many species are adapted to live in only certain environments.

One would not expect, therefore, to find Lepidosiren in the Alto Rio

Parana, the coastwise streams of eastern Brazil and Patagonia. Lepido-

siren lives in the vast swamps (chacos and pantanals) of Rio Paraguay

and Rio Amazonas.

4, Species of more or less recent marine origin have encountered far

greater difficulty in entering small, rapid, rocky, shallow streams with a

limited supply of food than large rivers, because marine fishes are used

to swimming in the sea and not in rapid rivers and because the change

from sea to fresh water is less sudden.

5. Some of these rivers have higher altitudes, and the number of fish

always bear a relation to the altitude.

The last of the environmental complexes which needs a further con-

sideration is that of Patagonia. It lies between 40° and 55° south

latitude and is characterized by a general paucity of plants and animals,

especially tropical forms. Its plains are arid and forestless. Its rivers

are not large, because they are for the most part fed by the melting

snow on the lofty Andes. A cold Antarctic current flows along the coast,

which is devoid of swamps. Besides, vast tracts of Patagonia have been

under the sea during part of the Tertiary period. At this time, it is

quite possible that nearly all of its fresh-water life was exterminated,

and as the land rose again from the sea with the Tertiary elevation of

the Andes, the northern portion of it became semi-desert. In part of

this region, the rivers flowing down from the Andes dry up on the barren

plains. Hence, a southern migration of fishes would have been almost

blocked, excepting in the case of the Pygide, which are found everywhere

in South America.

Notwithstanding all this and the fact that the Patagonian rivers have

few marine immigrants, its twenty-six known species of fishes contrast

favorably at least in number, either with the secca (dried) regions of

Ceara and Pernambuco, Brazil, or with similar latitudes in some parts

of the northern hemisphere. It is not at all strange that such tropical

74 ANNALS NEW YORK ACADEMY OF SCIENCES

species of fishes as Arapaima, Osteoglossum and Hlectrophorus are not

found in Patagonia, because they are also not found in La Plata; but it

is strange that the Pygide are found in Patagonia and Hoplias mala-

bricus (of the Characinide) is not, because I found Hoplias to be one of

the best overland travelers of all the South American fishes. The absence

of Hoplias in Patagonia may be due to its being a tropical genus.

The fact that Geotria and the Galaride are found in the Australian

realm is no evidence that Patagonia was connected with the same, be-

cause at least one of these forms is known to enter the sea.*® The

absence of Diplomyste and Pygide from the Antarctic and the Aus-

tralian realms seems to me to be far more conclusive static evidence that

these regions were not continuous than do the presence of Geotria and

Galaxide in these two regions indicate that they were continuous. The

latter two genera could have extended their limits of distribution by way

of the sea. If a connection existed, Pygide, being good overland tray-

elers, would have had a chance to enter the Australian realm. If Diplo-

myste is the most primitive living catfish, it, too, would have had a

chance to extend its limits of distribution. Hence, only marine fresh

water and no strictly fresh-water species or genera are common to these

regions, and I take this as strong evidence against a former connection

between Patagonia and the Australian realm. In fact, Patagonia has no

Osteoglosside, no Dipnoi and other forms found in Australia.

Origin of the South American Fishes

In the Princeton Patagonian report on the fishes, Professor Higen-

mann states that fishes probably interchanged before the beginning of

the Tertiary epoch between Africa and South America by way of a land-

bridge between Guiana and Africa. The following objections can be

raised against this hypothetical view:

1. There is no positive evidence that either the Characinide or the

Cichlide as such existed previous to the Tertiary, but I grant the possi-

bility of their existence in late Cretaceous times.

2. All of the known fossil fishes indicate a northern origin of the living

tropical fishes.

3. There is no good geological evidence in favor of the connection.

The evidence is all biological and paleontological and questionable in

kind.

%>It is also probable that formerly both of these genera were able to enter the sea.

This is all the more true in view of the fact that much of Patagonia was covered by

Tertiary sea. ;

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA %5

4, The alleged support derived from the distribution of the fishes is

derived from the static viewpoint: of animal geography. It is also based

on several erroneous ideas concerning the topography, geology and en-

vironmental complexes.

5. The point of family origin was not correctly determined, because

the greater number of species in a given locality is no evidence that it is

a center of family origin or dispersal.

In the preceding pages, I have attempted to explain the present dis-

tribution of fishes as being primarily due to cenotelic changes produced

by different environmental complexes on the ancestral stock. It was also

noted that only paleotelic characters were widely distributed. Therefore

we must look for the point of family origin at a very remote epoch, when

few or none of living Cichlid and Characinide existed. In other words,

the present distribution has little or nothing to do with the point of

family origin. This being the case, we have to look for fossils.

We must confess that no absolutely conclusive knowledge derived from

either paleontology or any other source exists from which we can defi-

nitely determine the point of origin of the cichlid and characinid fishes ;

but the analogies which can be drawn from the following data taken in

connection with the facts which I have already stated appear to give the

most plausible explanation, because it is more in harmony with the known

geological data.

The distribution of living and extinct Osteoglosside is as follows:

Number of

spectes Remarks

Phareodus (Leidy)....... 2 Eocene of Green River, Wyoming, United

States of America, and upper Cretace-

ous of Chico formation of western

United States of America.

PS RUCIO@UUSE. oc cris alone oes 1 Lower Hocene of England.

PAGE DOIG, Jo.0.0.s = elise 0 6 ere eres 1 Amazon and Guiana.

Osteoglossum ............ 3 Amazon to north, East Indies and Aus-

tralia.

ERCLCTOUISS Aco ais vise) cscs 5 0.0 il Tropical Africa.

Besides these five genera, two more doubtful genera are known from

the late Mesozoic of the United States of America and southeastern Eng-

land. The above data indicate a northern origin for this family of fishes,

which now lives only in the tropics and the southern hemisphere.

Diplomystus, a clupeoid, has recently been divided into two genera by

Jordan, and at least seven species are known. They are from the fol-

lowing regions:

7T—NY

76 ANNALS NEW YORK ACADEMY OF SCIENCES

Eocene—Green River, Wyoming, United States of America.

Upper Cretaceous—Mt. Lebanon, Asta.

Cretaceous—Mediterranean Islands.

Upper Cretaceous—Bahia, Brazil.

Lower Oligocene—Isle of Wight, England.

Upper Cretaceous—Italy.

Living forms closely related to these fossil species are said to exist in

Chili and New South Wales.

Priscacara, with seven species from Green River and Bridger Hocene

of Wyoming and Utah are so far the only known fossil Cichlide.** This

indicates a northern origin for the Cichlid. One genus with three spe-

cies of fossil Pomacentride are known from the upper Eocene and lower

Miocene of Italy.

One species of Perichthys is known from Tertiary shales of Taubate,

Brazil. This genus still lives in Patagonia and Chili.

At least seven genera with forty-three species of fossil Labridz are

known from the lower Eocene to the lower Pliocene of Europe and Hng-

land and the Eocene and Miocene of New Jersey, United States of

America. One tooth attributed to a member of this family is known

from the late Tertiary of the Argentine Republic.

About fourteen genera of Cyprinide, including about thirty-three fossil

species, have been reported from Germany, Bohemia, Sumatra, Java and

various parts of western United States of America. These fossil forms

range from the Quaternary to lower Miocene (?). Living forms are found

all over the world, excepting Australia and South America. The absence

of the Cyprinide from South America is the most extraordinary fact in

the distribution of fishes. They have existed since the Miocene in south-

western Idaho and are now found in Mexico but not in South America.

Furthermore, if an Archhelenic land-bridge existed between Africa and

Guiana and this was the means of dispersal of Characinide and Cichlidex,

why did not the Cyprinidz also enter South America? Tetragonopterus

avis and T. lignitis .(Hobrycon Jordan) from Taubate, Brazil, from

shales of doubtful late Tertiary age are so far the only definitely known

fossil Characinide.

_ 86 Woodward, 1898, reported a fragment from Taubate as Alquidens (?).

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA “%?

The facts regarding the Siluride are as follows:

Number of

species Remarks

\CUTTATIS 9 5a nk ee 1 Lower Pliocene, India ; still living in India.

Heterobranchus .......... 1 Lower Pliocene, India; still living in Africa

and East Indian Archipelago.

ST (?) Upper Tertiary of Europe and India; still

living in Paleoarctic realm.

Pseudeutropius .......... il Tertiary of Sumatra.

Macrones ............+.+. 1 —_ Lower Pliocene of India; existing in Asia.

BUD vere erence cece eeeeee 1 Lower Pliocene of India; existing in Asia.

AMIUTUS .0 0s esseeeerecees 2(?) Lower Miocene of Canada; existing in

.North America and China.

(?) Pimelodus ........... 1(?) Late Tertiary (?) of Taubate lignite,

Taubate, Brazil, and of Parana River,

Argentine Republic; existing in South

America east of the Andes and north of

Patagonia.

BGIUVIVEUSECSI. or <5 0\0) 0: 61 sis's 010-5 7(?) Lower Tertiary, lower Miocene and

Bridger Eocene of western United States

of America.

Bucklandiwm ......es.00% 1 Lower Hocene, England.

ALONG (5 OSA 4(?) Middle Eocene, Belgium.

Upper Eocene, England.

Oligocene and Middle Oligocene, Germany.

Lower Hocene, Copenhagen.

Pliocene, India.

(?) Tertiary, Taubate, Brazil.

RUG QGUUS) che o04 6.6 010 siosisiee's 1 Tertiary of Sumatra.

Woodward stated that Bucklandium diluvii appeared to be related to

Auchenoglanis, which still exists in Brazil. Rhineastes is the oldest

known fossil catfish and appears to be related to Phractocephalus of the

Pimelodine, which lives in the Amazon.

The age of the Taubate shales, found in the deeply eroded Parahyba

Valley, is not definitely known, but the surface deposits containing fishes

do not appear to be very old (Pliocene), because the three fossil genera

of fishes are still living. Jordan’s generic distinction of Hobrycon does

not appear to the writer to be well founded, because the species of both

Astynax and Tetragonopterus vary considerably in shape. The two fossil

species from Taubate may, in fact, fall into the genus Astynax as now

defined by Higenmann.

If the writer is correct in considering the Taubate shales as late Ter-

tiary, it is evident from the above list of fossils that the South American

as well as the African fishes have evolved from forms which earlier lived

78 ANNALS NEW. YORK ACADEMY OF SCIENCES

in the northern hemisphere. In fact, Osborn has already stated in “The

Age of Mammals” that the South American fishes show an ancient north-

ern affinity. The dipnoans and crossopterygians, which are now found

only in the southern hemisphere, also were northern in origin, as far as

we now know.

The fact that South America has many species but few families of

fishes also vaguely indicates a northern origin, because both plants and

animals often rapidly break up into new species when placed in new

environments. If the Cichlid were not northern in origin, how were

they able to get into Wyoming during the early Hocene, when they are

now not able to get north of Rio Grande and have never been able to

enter Patagonia? The mere fact that the characinids and cichlids are

still “going wild” in making species indicates that they entered South

America during or after the late Miocene, 7. e., after South America be-

came permanently connected with North America. This view seems to

be all the more probable, because it appears to be easier for animals to

move from temperate regions to the tropics than vice versa.

It must also be noted, before dismissing the subject of the point of

origin of the South American fresh-water fishes, that there is some vague

evidence in favor of the marine distribution of at least the Cichlide and

Osteoglosside. The genus Priscacara is closely related to the marine

Pomacentride, according to Cope, and the formation in which they are

found appears to have been near the sea level. Hence, in view of the

fact that the actual paleotelic or ancestral forms which were distributed

are not definitely known, a marine distribution of many primitive forms

is not at all impossible. Such a view is made all the more probable by

actual experiment as already noted.

The present distribution of the osteoglosside can be explained most

easily by considering them as northern in origin, at least if the zodgeog-

raphers do not entirely ignore the paleontological evidence. It must be

admitted that there is no positive evidence to show that the Characinide

are directly northern in origin, but the only positive evidence at present

known indicates that both the Cyprinide and Nematognathi, the nearest

relatives of the Characinide, originated in the northern hemisphere, and

it is highly probable that the ancestral Ostariophysi were also northern

in origin—if we admit that the Ostariophysi are a homogeneous group.

In this connection, the Nematognathi appear to have first split off from

the ancestral Ostariophysi. Then the Cyprinide split off, and some of

the later Ostariophysi were pushed, after the Miocene times, into South

America, and the Characinide, now found in the southern hemisphere,

appear to be the lineal descendants of this ostariophysian stock and have

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA %9

therefore retained more of the primitive characters than either the

Nematognathi or the Cyprinide.

There have been sufficient connections between North America and

South America and between Eurasia and Africa to permit exchange of

fishes during past epochs. Besides, many fishes and other fresh-water

forms may have been able to migrate short distances along the coast or

from island to near-by islands. Hence no objections can be raised against

such migrations for want of land connections.

SUMMARY OF THE MOST IMPORTANT DATA WHICH HAVE BEEN USED TO

SUPPORT THE VIEW THAT SouTtH AMERICA AND THE HASTERN HEMI-

SPHERE WERE PRIMITIVELY CONNECTED.

INVERTEBRATES

Crustacea

Ortmann has stated that only one family of crabs found in northern

South America lends positive support to his view of Archhelenis, 7. ¢., a

pre-Tertiary connection between Guiana and Africa. This family, Poto-

mocarcinine, are found only as far south as Guiana. They are found in

Central America. Why have they not immigrated into the Amazon Val-

ley and south? For all we know, they may have immigrated from the

northern hemisphere and have only reached Guiana.

Their ancestral stock might have been drifted across the Atlantic by

the African-West Indian current or their ancestral stock, which was dis-

tributed, may easily have been distributed by the way of Europe and

North America. This is especially true in view of the fact that no iden-

tical forms are found in Africa and South America. What was this

paleotelic form which became distributed and gave rise to the different

genera of the eastern and western hemispheres, and where did it origi-

nate? This family of crabs offers a splendid analogy to the present dis-

tribution of camels and tapirs which now live in the tropics, but which

were originally found in the northern hemisphere. Besides, the Mesozoic

forms of crustacea are very imperfectly known, and Sd is especially true

of the crabs in question.

Mollusca

The writer found a bivalve, Diplodon, in the Iguasst. River above the

big falls, which in some form or other appears to date back to the Tri-

assic. This does not necessarily mean that Diplodon dates directly back

to the Trias, because this genus is widely distributed over the highlands,

but it does mean that it is a primitive form. In contrast to the above

80 ANNALS NEW YORK ACADEMY OF SCIENCES

genus is Hyria, which was seen by the writer only in the lower Amazon

and therefore probably does not belong to the older highland stock. I

did not see a dozen species of bivalves in Rio Guaporé, which probably

indicates a:rather primitive stock, but they were very abundant in Rio

Uruguay, which indicates a great cenogenic evolution in this region.

In a general way, the mollusca follow the same rules of distribution as

the fishes, but our knowledge, especially from accurate field data, of the

fresh-water bivalves of South America is entirely too meager to be used

in support of any theory. For example, Ortmann has found that some

of the bivalves have palpi, arrangement of gills and siphonal openings

like some of the African bivalves, but this may be due in both cases to

living in muddy, tropical water. Before such evidence can safely be used

to support any theory, much careful field work and experimental evi-

dence is needed. That is to say, we need dynamic and not static data.

Many of the fossil bivalve shells in North America resemble living

forms in South America. Inasmuch as the soft parts are not preserved,

it will be very difficult to determine the paleotelic forms of bivalves which

were distributed. Von Ihering has listed 581 species of mollusca along

the coast of Brazil, 54 of which are known on both the Antillean and

African coasts, while 72 are found in common on the African and Bra-

zilian coasts. He does not appear to attach much importance to larval

distribution of these forms, but it is well known that some larval crus-

tacea and mollusca are found on the high seas. Marine turtles also cross

the sea. Nichols has recently reported a case in which he thinks that the

mid-ocean whirlpool and side currents swept young T’rachurus from the

coast of England to the coast of Florida.. These ocean currents are well

known to the sailors, who often go far out of a direct route in order to

avoid them. In some places, when the winds are favorable, the one off

the Barbados Islands is said to run about five miles per hour. Currents,

therefore, may have transferred some mollusca between the South Amer-

ican and African coast along with sea weeds and other drift. Then,

again, a few forms could have been transferred by primitive sailing

vessels between the various ports.

It is interesting to note in this connection that practically all of the

mollusca known from the Brazilian coast are reported from near larger

or smaller seaports. In other words, very little of the Brazilian coast has

been surveyed, and until regions remote from seaports are carefully

studied, too much stress must not be laid on the present list of mollusca.

Besides, the above lists of common forms on the African and Brazilian

coasts include such widely distributed (cosmopolitan) forms as Mytilus,

which could easily have gone south from the European and North Amer-

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 81

ican coasts. The writer looked carefully along the coast south of Iguape

for Aporrhais pespelecani and was unable to find this form either living

or dead. The few dead shells of this species known from a seaport have

_ little significance, because peddlers and sailors are known to be great dis-

tributers of shells. The writer picked up one valve of Lucina jamai-

censis (?) on a sand bar below the Urubu-punga waterfalls of the Alto

Rio Parana, which is several hundred miles from the seacoast, but this

shell had evidently been dropped there by an Indian.

A very important factor in the distribution of the marine mollusca of

the Atlantic Ocean is the tropical condition which existed in the North

Atlantic during the Eocene. This would have given excellent oppor-

tunities for exchanges of forms between the African-Huropean and

American coasts. These ancestors of the existing forms would have been

pushed south again when the climate of the North Atlantic became

cooler. As a result of this, many resistant ancestral forms living in

similar environments and evolving along rather definite lines would pro-

duce a great similarity between the coasts of the South Atlantic.

Furthermore, it is not impossible that some young forms of land

gastropods could have been carried with tropical plants to the eastern

hemisphere just as Litorina litorea has probably been imported in some

way to the American coast.

After a long detailed study of the living and fossil Tertiary mollusca,

von Ihering has recently concluded that Archhelenis, the land-bridge

between Africa and South America, began to disappear in the Cretaceous

but continued to exist in the Tertiary. Ortmann (1910) used the same.

data and arrived at a different conclusion, namely, that Archhelenis had

disappeared before the beginning of the Tertiary; but neither of these

authors has taken into consideration the effects of similar tropical en-

vironmental complexes along the African and South American coasts on

the ancestral stock from which the existing species have evolved. When

this is done and the cosmopolitan forms are eliminated and when due

allowance is given possible larval and adult distribution by ocean cur-

rents, floating debris and boats, then no land-bridges are needed to ex-

plain the distribution of marine mollusca.

Only static studies have been made, and until some » dynamic work has

been done the evidence derived from the mollusca is not a safe peg to

hang a theory on.*”

87 The brachiopoda appear to me to offer even less evidence, because many almost cos-

mopolitan genera have existed during various past ages. This is all the more true when

certain similar forms are known to exist in similar environments. So it appears that

extensive land-bridges like Archhelenis will have to rest on evidence derived from geology

and continental faunas and floras.

82 ANNALS NEW YORK ACADEMY OF SCIENCES

Recently Pilsbry has given an excellent static treatise of the distribu-

tion of non-marine mollusca of South America. He has combatted the

separation of Guiana, Brazil and Patagonia during past times. In this

far, I agree with his conclusions, but his data do not alone warrant them,

because the highland environments in Guiana and Brazil are very much

alike, and hence similar forms are to be expected from a common older

stock, even if these regions had been isolated from each other during part

of the Mesozoic and Tertiary times. His static evidence certainly almost

if not completely destroys that of Ortmann for a Guiano-African con-

nection, but he has unfortunately located his Brazilian-South African

connection exactly in the region where we have given extremely strong

geological evidence against such a view. ‘The entire coast of Brazil in -

this region is fringed by marine Cretaceous, which alone would force this

connection to have disappeared at least in the early Cretaceous. Besides,

the “Pernambucan fan” is strong evidence against any Paleozoic connec-

tion in this region. ‘There is also so much other geological evidence against

the building up of a land-mass across the great ocean depths of the South

Atlantic that we may consider Dr. Pilsbry’s view highly improbable, at

least until some dynamic and more careful field studies have been made

on the non-marine mollusca of the regions in question. His actual paleo-

telic forms which were distributed are not yet absolutely known to have

originated in this purely hypothetical Gondwana Land.

Ants

Von Ihering has even used the distribution of the ants to prop up his

Archhelenis theory. He states that we still see the ingression of Bolivian

ants into Brazil. These ants are supposed to have come by the way of

Antarctica from the eastern hemisphere. This evidence appears to me to

be of little weight, because the ants have had a highland since the Per-

mian, over which they could have crawled or flown into Brazil. Further-

more, practically nothing is known of the ants which live in the high-

lands of central Matto Grosso. If one may hazard a guess, I would sug-

gest that a detailed study of the ants would show an invasion from Brazil

toward the Andes and not vice versa, after the elevation of the mountains

in Tertiary times.

Corals

Gregory has shown that the Miocene corals of the Mediterranean re-

semble the living corals of the West Indies. This is not, I believe, evi-

dence that the West Indies were directly connected with the eastern

hemisphere, because the larvee of these corals may have drifted over by

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 83

the mid-Atlantic whirlpool and ocean currents. They may have origi-

nally migrated down the North American coast from Europe, and it is

- even more probable that the observed affinities are due to similar evolu-

tion in similar environments. If this is not true, why have so many

species remained similar to the Miocene forms of Europe ever since the

supposed land-bridge disappeared or from before Tertiary times? When

and where did this bridge exist? Are not the corals of the North and

South American coasts also similar? If not, why not??*

GONDWANA FLORA

Similar deposits of sandstones, clays, shales and bowlder till are found

in Brazil, South Africa, India and Australia. These deposits are chiefly

‘Permian and contain among other fossils the characteristic lower Gond-

wana flora (Gangamopteris or Glossopteris flora). The identity of the

Gangamopteris flora, along with many fossil and living genetically re-

lated animals found in Africa and South America, has led to a widely

accepted belief that these continents were originally connected. This old,

land-mass has been designated Gondwana and is thought by some to have

extended across the Atlantic between either Brazil or Guiana and Africa.

Others have ignored this connection and have maintained a southern

connection by way of the Antarctic Islands.

The Gangamopteris flora, according to I. C. White and David White,

has been found six meters above the crystalline floor of the coal fields of

southern Brazil. At this level, only Gangamopteris obovata was found.

The next higher level, 55 meters above the granite floor near Minas,

Santa Catharina, contains Rosellinites gangamopteridis, Hysterites bra-

stliensis, Phyllotheca griesbachi, P. mulleriana, Glossopteris browniana,

Werte0raria ........ ?..., Gangamopteris obovata, Arberia minasica,

Derbyella aurita, Noeggerathiopsis hislopi, Cardiocarpon seixas and

Cardiocarpon moreiranum.

These species belong to the early typical Gangamopteris or lower

Gondwana flora. The same genera, and in many cases identical species,

are found in the Ecca shales of South Africa, in the coal associated

with marine lower Permian of New South Wales and Tasmania and in

the Karharbari beds of India. The same flora is found in the lower

Coal Measures of Argentina and the Falkland Islands. Only much later,

in the upper Permian of the northern part of Russia, are any of these

*87f Archhelenis existed, I fail to see how the rivers could have been arranged on it so

that only one family of crabs, two families of fishes and a few fresh-water and land

mollusca took advantage of it, when the same theory assumes that the coastwise streams

of eastern Brazil have been barriers to at least part of this fauna.

84 ANNALS NEW YORK ACADEMY OF SCIENCES

gangamopterids known to occur in the northern portion of the northern

hemisphere.

Conformably underlying this lower Gondwana flora are the Orleans

conglomerates of southern Brazil, which are supposed to be related to the

Dwyka conglomerates of South Africa, the Baccas Marsh conglomerates

and their equivalents in Australia and Tasmania and the Talchir con-

glomerates of India. These conglomerates contain some of the evidence

for the alleged Permian glaciation of the southern hemisphere.

At 135 meters above the granite, 7. e., about 80 meters above the typi-

eal lower Gondwana flora, is found the intermingling of this flora with

some species of the older northern cosmopolitan flora. In this forma-

tion were found Hquisetes calamitinoids, Schizoneura, Sigilaria aus-

tralia, Sphenopters hastata (?), Glossopterts indica, G. ampla, G. occt-

dentalis, Neggerathiopsis hislopt and Cardiocarpon oliwveiranum. The

flora is still primarily Gondwana. At a still higher level, 157 meters

above the granite floor or 100 meters below the Iraty shales containing

Mesosaurus, are found more of the northern flora, such as Lepidodendron

perdroanum, Lepidophloios larcinus and Sigillaria brardw. At this

level, the lycopods are again preéminent as coal makers.

The Gangamopteris flora is very imperfectly known, but what is known

indicates almost beyond a doubt that the Gangamopteris belong to the

southern hemisphere. It is not known from North America and is only

known from the late Permian of Russia. The question is, then, Are the

known facts concerning the Gangamopteris flora indicative of a contin-

uous Gondwana Land somewhere in the southern hemisphere? Before

attempting to settle this difficult question, it is necessary to consider the

origin and the environmental complexes of this flora.*®

The Gangamopteris flora belongs, as Professor Arber and Dr. White

have well shown, almost exclusively to families already known in the

cosmopolitan flora. They constitute genera and species more or less

bound to their northern relatives, though often differmg much in form

and aspect. In general, they appear simpler in figure, with a tendency

to thickness and rugosity of leaves, and on the whole their general aspect

suggests environmental conditions unfavorable to luxuriant growth. . This

flora suddenly appears in the early Permian well defined from its Car-

boniferous ancestors, which lived in the northern hemisphere and sur-

vived more or less the profound geological changes produced by the for-

®9It is barely possible that the lower Gondwana flora of Brazil belongs to a later Per-

mian than now believed, as Professor Branner states in his Geologia Hlementar that the

intercalated marine deposits containing Schizodus, Myalina and Oonocardium also con-

tain other lamellibranchs also found in the Triassic. If this is found to be the case,

then the appearance of this flora in Russia-Siberia may have been as early as in Brazil.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 85

mation of the Brazilian Plano Alto. This cosmopolitan flora continued

to exist in the Mesozoic epoch, but the Gangamopteris flora, excepting

Glossopteris and Schizoneura, vanished with the close of the Permian.

In this remarkable fact, we have, I believe, part of the solution of the

Gangamopteris flora, 7. ¢., it existed in Brazil only during the formation

of the Plano Alto and died out after this was completed (early Triassic).

In this profound change of the ancient Brazilian topography produced

by the formation of the Plano Alto in the Permian inland basin, we have

the production of unfavorable environments which might have produced

the alleged glacial effects on the Gangamopteris flora and have caused

the absence of the cosmopolitan plants in the lower Gondwana formation

of Brazil. This view is further strengthened by the fact that the shales,

etc., of the lower Gondwana formation have a different appearance and

chemical analysis from those of the higher formations in which the cos-

mopolitan flora are found. The cosmopolitan flora is always associated

with the production of large coal fields, and such conditions are not met

in the Gondwana formations of Brazil. It appears to me that it was

desiccation (perhaps not due to a lack of rainfall, but to its disappear-

ance in a sandy soil) and the blowing of sand into the Permian inland

basin, and not severity of climate (glacial), which produced the stunted

appearance of the early Gondwana flora.

Several other objections can be raised against the use of Permian

glaciation as a factor which affected the distribution of the Permian

plants and reptiles of South America. In the first place, the writer does

not believe that the existence of glaciers in Brazil has been definitely

established. His experience with glaciation in North America and gla-

ciers in the Andes, taken in connection with observations on erosion in

the highlands and mountains of Brazil, has strongly suggested to him

that the Orleans conglomerates were not deposited by glaciers. In the

highlands of Piauhy and various places in Brazil, one can see both high-

land streams and extensive slanting surfaces over which gravel and

bowlders slide during heavy rains. The underlying surfaces and bowlders

are often scratched in a way which resembles glaciation. When pieces of

the scratched and polished surfaces are detached, segregated or not, as is

often the case due to less and greater amount of rainfall, and deposited

at a lower level, a “false moraine” and even false bowlder till is formed.

When such a mass of gravel, clay, bowlders, etc., becomes covered up and

pressed together by later erosion in little stratified or unstratified beds

(due to continual deposition and plasticity of the clay), it can easily be

mistaken for a glacial deposit. It can only be distinguished from glacial

deposits by means of truly faceted bowlders; and inasmuch as faceted

86 ANNALS NEW YORK ACADEMY OF SCIENCES

bowlders are not definitely known to exist in Brazil, I take this as a

strong evidence against Permian glaciation in Brazil. In fact, not until

recently has any one even seen striation in the alleged glacial deposits of

the Permian of Brazil. Some of these recent false signs of glaciation in

Brazil, as Branner has shown, even deceived Agassiz (the expounder of

glaciation), who described vast sections of Brazil as being glaciated.

The direction of the striations and the arrangement of the bowlders

also offer no conclusive evidence in favor of glaciation. When “false

moraine,” composed of scratched surfaces, bowlders and false tillite, be-

came covered up by the Permian sandstone found in the Plana Alto,

only those deposits and scratches pointing in the direction of the dip of

the country were exposed by the post-Permian erosion of the overlying

strata. In most of the Gondwana formation, this dip is toward the south

and west. Hence only here and there are the strie exposed, and all of

them point more or less in the same direction, 1. e., they are only seen

along deeply eroded river valleys below the waterfalls. Furthermore,

these deposits often cover vast regions. For example, the deposits of

such erosion would have covered many miles of width for the entire

length of the Serra do Mar during the late Carboniferous and early Per-

mian when the climate was favorable for the deposition of false bowlder

tillite, which later became overlapped by the Permian sandstone. Later

erosion exposes these deposits over a vast area.

Woodworth found striations on bowlders, some of which appeared to

have been deposited by ice floating near sea level (as is indicated by in-

tercalated marine deposits in the Rio Negro basin). This floating ice

may have come from Permian swamps, where it gathered up bowlders.

This mass may have floated toward the sea the following season, scratch-

ing the rocky surfaces along the margins of the swamps. In this case,

faceted bowlders would probably not have been formed. I acknowledge

that Woodworth has also found much other evidence for Permian glacia-

tion in Parana, Brazil, and the evidence is even stronger for it in the

eastern hemisphere.*°

I grant that glaciers may have existed in Brazil during the lower Per-

mian epoch, but in view of the preceding, it appears that faceted bowl-

ders must be found before the evidence in favor of it is sufficient to

warrant the use of Permian glaciers as a factor in the distribution of

plants and animals. Even if such evidence is found in Brazil, it will

probably not be found after the lower Permian, at which time the typical

Gondwana flora and reptiles were scarce. Furthermore, if glaciers ex-

40 Woodworth had not published his paper on the Permian glaciation in southern Brazil

when these notes were prepared.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 87

isted during the lower Permian in Brazil, they may also have been only

local glaciers of high altitude, like those existing in the Andes.

Returning again to the fact that the Gangamopteris flora only existed

during the formation of the Plano Alto, it appears to me that this is

positive evidence that its environment, composition and extinction were

directly associated with this great transformation of the South American

topography. In other words, the Gondwana flora was an arid highland

flora.

It is known in marine series in Australia, but it could have been washed

there from higher elevations, and the altitude of these sandy highlands

was not necessarily great, in view of the fact that stunted plants are

found on the Brazilian highlands at comparatively low altitudes. Similar

geological transformations also occurred in India, Africa and Australia.

Similar deposits of shales, clays, sandstones, coal, etc., occur in these

remote regions. ‘The mere fact that all of these regions possessed almost

identical environmental complexes, not known to be perfectly duplicated

in the northern hemisphere, is one of the most important factors con-

nected with the origin, the distribution and extinction of the Gangamop-

teris. flora, regardless of whether these regions were or were not con-

tinuous.

It is a remarkable fact that so many identical species and genera

belonging to different families of both the Gangamopteris and the cosmo-

politan flora could exist, unchanged, in such remote regions as India

and Brazil during most of the Permian epoch. Little short of ortho-

genesis or similar evolution in similar environments can account for such

data.

I have already shown that the Permian inland basin of South America

is almost completely surrounded by higher Archean mountains which

have apparently remained almost stationary during part of and since the

Permian epoch. This is particularly true of southern Brazil. There-

fore, the fact that the Gondwana flora, so widely distributed over the

southern hemisphere, was able to enter these various Permian formations

over and around higher Archean mountains is evidence that its ancestors

- were little or not at all affected by barriers. It is true that this flora

might have entered the outlets of the basin along the Permian coasts,

but if it was a highland flora, it probably did not.

I am unable to conceive how and from where enough of the typical

deposits of Gondwana Land could be derived in such a way that a

homogeneous environment might have existed between either Africa and

South America, or between South America by way of the Antarctic

islands and either Africa or Australia. The Gondwana formation of

88 ANNALS NEW YORK ACADEMY OF SCIENCES

South America can easily be explained by place erosion of the highland

pre-Permian floor, by the erosion of the Serra do Mar and its southern

spur, Serra Geral, on the east side of the Permian inland basin, and by

similar changes of the Serras de Cordova, San Luis and de la Ventana

on the west. All these mountains have a general north and south trend,

and there is no evidence of an east-west trend in any of this region. The

Gondwana formations of Brazil, Africa and India dip as a whole toward

the south and west. This has a ready explanation from the location of

the Archean and pre-Permian rock, but there appears to be no explana-

tion showing how these formations could have been a part of a greater

continuous homogeneous Gondwana Land. Its flora being only known

from this special type of environment, how did it traverse these obvious

Gondwana barriers ?

Permian and Carboniferous deposits of coal are laid down along the

sides of the Andes-Rocky Mountain system which extends into Hurasia

and also on the sides of the Appalachian system, and extending through

Guiana down the divide along the eastern coast of Brazil. I am not

aware of any similar formations extending through the Antarctic islands

toward either Australia or Africa. The Antarctic islands appear to bear

the same relation to southern South America as do the West Indies to

northern South America. They are well separated from both Australia

and Africa by great depths of the ocean as well as by many miles of dis-

tance. Hence it appears more probable that the Gondwana flora or its

ancestral forms would have migrated along lines where the conditions of

the environments of Gondwana Land were at least partially duplicated,

1. €., Where coal was deposited, rather than over about 2,000 or more

miles where such conditions probably did not exist.

The fact that the northern Permian plants had many identical species

in the southern hemisphere, that there is considerable difference between

the Permian reptiles found in Texas and those of South Africa, as well

as the fact that the Gangamopteris flora is found in the late Permian of

Russia, is good evidence that both the cosmopolitan and the Ganga-

mopteris flora were resistent forms which (or their ancestral stock) could

migrate into remote regions with comparative ease. The presence of ~

Cardiocarpon, Sphenopteris, Psaronius, Sigillaria and Lepidodendron in

both North America and South America, as well as the presence of the

Mississippian flora at Cacheuta, Argentina, indicates an exchange of

plants between North and South America. Some of these plants (Lepi-

dodendron) are not yet positively known in South Africa. The cosmo-

politan flora is also known from Europe and other parts of the eastern

hemisphere, and it is remarkable that this flora could not only become so

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 89

widely distributed but could also remain almost identical in the various

corners of the earth.

Hyen if it be granted that a vast homogeneous Gondwana Land en-

tirely covered the southern hemisphere, this alone would not explain why

so many species of the typical lower Gondwana flora remained identical

during the greater part of the Permian epoch. There is positive evidence

from the intercalated Permian of Brazil and other Gondwana forma-

tions that these regions were not continuous during the entire Permian

epoch, and yet the flora remained identical. Hence there appears to be

no need for a ready and wholesale exchange of this flora, because it re-

mained unchanged during the most of the Permian epoch, and therefore

any accidental distribution of each genus and for one time only would be

adequate. In this connection, it is to be remembered that neither con-

tinuous land connections nor barriers nor isolation causes species (unless

there are only a few individuals) to evolve new forms or to remain fixed,

but the action of the environmental complexes on the ancestral forms,

which had a certain composition, and hybridization will change the spe- .

cies. The identity of the Gangamopteris flora in such remote parts of

_the earth as already noted may be due to the action of similar environ-

mental complexes upon the ancestral stock of each group of this flora.

It is then even possible, but not probable, that the Gangamopteris flora

is only an environmentally changed form of the cosmopolitan flora which

could not exist as such in the early Gondwana environment. At least the

sporaginous members of the cosmopolitan flora could have migrated into

the Gondwana environments, if they had been able to thrive under such

conditions.

What the direct antecedent types of the different groups of the Gond-

wana flora were is not definitely known. David White considers that

Neggerathiopis is probably of Cordaitalean origin and Gangamopteris

has a common origin with the neuropterid group of Cycadofilices, to

which the genus Glossopteris also is related; yet both are far removed

from the known antecedent type of the northern hemisphere. What this

direct ancestral stock was and where it originated are questions which

must be answered before we can definitely hypothesize lines of dispersion.

I have repeatedly pointed out in the case of fishes that direct connec-

tions are not necessary for the production of identical species and that

the greater number of species in a given region is no evidence that the

family originated there. I will repeat here again that the fact that the

gangamopterids are found only in the southern hemisphere is not con-

clusive evidence that their ancestral stock either originated there or that

all of the existing Gondwana formations were directly continuous.

90 ANNALS NEW. YORK ACADEMY OF SCIENCES

The Gangamopteris flora was already well defined in the early Per-

mian. It has no close antecedent types in either the northern or the

southern hemisphere, but its actual origin must have antedated the

oldest formations where it is known to exist. It may well have been

these ancestral types (like the highland fishes of Brazil) which were dis-

tributed and not each individual species. These antecedent types may

have arisen from the northern cosmopolitan flora and later have migrated

into the southern hemisphere where this ancestral stock of the different

groups underwent similar evolution in the similar environments of the

Gondwana Land. This view is supported by the fact that these ancestral

forms would not have necessitated a homogeneous environment, because

they were more generalized forms.

Two objections may be raised against this view. First, it is conceiv-

able that Gangamopteris or various individual genera may have origi-

nated independently in different continents or in different parts of the

same continent, but the assumption that a flora of characteristic asso-

ciation and unity as the gangamopterid originated in this way is highly

improbable. It is not so improbable, however, if we include representa-

tive types of each group in the distributed stock and if we grant, as the

facts indicate, that identical evolution took place in similar environments.

Secondly, there is no fossil record of the gangamopterids in North

America. This objection to either a northern origin or a distribution of

the ancestral genera by the way of North America is, I believe, far from

being a fatal one. If the progenitors of the gangamopterids passed

through North America, the line of migration may have been along the

Appalachian system during the Carboniferous epoch, but the line of

migration was more probably from eastern Asia by way of Alaska. At

that time, the necessary Gondwana environment for the Gangamopteris

flora may have existed in North America-for a brief period. If it did

not, we only have to assume that the progenitors of this flora did not ~

necessitate Gondwana environments for a ready distribution. Dr. White

has told me that there is a little evidence in favor of a Cordilleran

migration, and recent work in the Permian of Texas indicates the same.

The flora would not have required a vast period of time to inter-

change between Eurasia by way of North America into South America.

The chances are that no trace of these typical Gondwana environments

or the ancestors of its flora would remain in either the Appalachian or

the Cordilleran regions, because they have been leveled to the sea, except-

ing in small patches, and re-elevated several times since that epoch. If

the flora was a highland flora, we now know where to look for it in

western North America.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 91

Tn this connection, great emphasis must be placed on the stability of

the Serra do Mar and the surrounding Gondwana formations of southern

Brazil when contrasted with the violent geological transformations which

the possible regions of migration through North America have under-

gone. In fact, I firmly believe that no traces of the Gondwana flora

would remain in southern Brazil, if this region had undergone the same

radical changes as the possible routes of migration through North

America.

The mere fact that the Gangamopteris flora appeared to live under

rather unfavorable conditions in which little coal was deposited is very

important. Could not similar Gondwana environments have existed dur-

ing a brief period in the northern hemisphere, and might not these en-

vironments have been obliterated by the post-Permian changes of the

North American topography? It appears that the stability of the Plana

Alto has saved the Gondwana flora of southern Brazil.

Professor Branner states in his “Geologia Elementar” that marine

fossils are found associated with the deposits containing Stereosternum

in the State of Sao Paulo. These fossils are all lamellibranchs, which

include such Permian genera as Schizodus, Myalina and Conocardium,

and other genera which are equally well considered as Triassic. This is

the only known invasion of the sea that entered the Gondwana region of

Brazil during the Permian and subsequent periods. Hence the Gond-

wana of Brazil has been extremely stable. This indicates very great

altitudes along the eastern side of South America during the Paleozoic

epoch. These great altitudes are needed: to build the great Plano Alto

and to account for the existing altitude of Serra do Mar after the vast

ages of post-Permian erosion.

If the Gangamopteris flora entered South America from North Amer-

ica, one may ask why it is not found in northern South America.

Whether it came from North America or not, I believe that it existed in

northern Brazil, because the Permian inland basin was continuous from

southern Brazil to the Guianas. In fact, Psaronius has recently been

found by Dr. Lisboa at Floriana on Rio Parahyba do Norte. All this

part of South America is very imperfectly known and awaits explora-

tion. If the Gondwana flora is later found in northern South America,

about 3,000 miles of a northern distribution will be abridged and the

remaining distance will not be many times greater than a possible south-

ern one. Hence due allowance must be made not only for the imperfec-

tion of the fossil record but also for the lack of sufficient exploration.

When this is done, a northern distribution of the Gondwana flora is not

altogether impossible.

S—NY

92 ANNALS NEW YORK ACADEMY OF SCIENCES

As an example of the imperfection of the fossil record may be given

Cryptobranchus, an amphibian, living in Japan and the Mississippi Val-

ley. The only known fossil relative is Andrias, Scheuzer’s Homo diluvii

testis of the Miocene of Europe. Its distribution, in a way, is like that

required for a northern dispersal of the Gondwana flora. As an example

of the imperfection of exploration may be given the two new marine

horizons in the Conemaugh series of western Pennsylvania, found by

Raymond in a region which has been explored by many geologists. As I

have already stated, however, there is little hope of finding the necessary

antecedent types of the Gangamopteris flora in North America, because

the necessary highland deposits which produced this stunted flora have

apparently disappeared, or have not yet been found.

The idea of a continuous Gondwana Land has little or no support

other than indecisive statistical data derived from the distribution of

living and extinct animals and plants. Furthermore, it appears to me

that even such a vast amount of indecisive data which admit of a variety

of interpretations can never outweigh the fact that the Permian reptiles,

the ooze and shark teeth, etc., of the great ocean depths, such as exist

between Africa, Australia and the Antarctic islands, the geology of the

Brazilian coast, the absence of giant east and west trend lines of South

America and the Antarctic islands offer strong positive evidence that a

continuous Gondwana Land did not exist. Until this vast array of posi-

tive evidence in favor of the persistence of the great ocean depths and

the continental shelves has been satisfactorily accounted for, it is just as

inconceivable to explain the existence of a continuous Gondwana Land

as to conceive either a northern distribution of the Gangamopteris flora

or that this flora developed orthogenetically from some unknown north-

ern ancestors which evolved from the Devonian cosmopolitan flora. This

is all the more true when we have to acknowledge that the actual point of

family origin of the Gangamopteris flora is unknown.

Before, however, the reader forms an opinion concerning the existence

of a continuous Gondwana Land, as is indicated by the distribution of

the Gondwana flora, he should consider still two other possible means of

distribution of the Permian plants which require no continuous Gond-

wana Land.

The distribution of the Gondwana flora by strong winds like those

which blow dust from the pampas of Argentina to Africa is possible but

perhaps not probable. The Gangamopteris plants are now generally

considered to be seed-bearing, but in most cases no seeds are known;

hence wind distribution would be out of the question, unless the seeds

were of very light-winged type. Then they might have been blown for

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 93

some distance, such as might have intervened between South Africa and

the nearest Antarctic islands. If any of them were spore-bearing, then

wind distribution would be an important factor, as it is in the case of

puff-balls and ferns. It is then important to settle definitely whether any

or all of the early Gangamopteris flora were seed-bearing or sporogenous.

The last possible mode of distribution is by the way of the sea. David

White has informed me that some of the seeds may have been able to

survive marine drift for some time. He thinks that the migration was

by the Antarctic, and if by the way of the sea, it would have been with a

minimum interruption by water. He suggests that it would perhaps be

better to say that migration was probably by several lands and not by a

continuous Gondwana Land. As the facts indicate, however, this flora

was a highland flora, and hence few or none of the species could have

been distributed in this way unless they lived on low coastal sandy high-

lands or campos such as exist in parts of Rio Grande do Sul, Brazil.

In conclusion, we may safely say that we do not definitely know where

the Gangamopteris flora originated, how and which way it dispersed,

why it appeared and disappeared in Brazil during the formation of the

Plano Alto and why so many species and genera remained almost iden-

tical in such remote regions as India and Brazil during most of the

Permian epoch.

The distribution of the Permian reptiles, the deep intervening sea, the

trend of the Archean mountains, the mode of the formation of the Per-

mian deposits, the location of marine deposits and the evidence in favor

of the persistence of the great ocean depths and the continental shelves

offer conclusive evidence that no continuous Gondwana Land has existed

between South America and the eastern hemisphere, at least since Car-

boniferous time. Previous to this, it may have existed, but many data

are needed to prove that it did.

In view of the fact that the Gangamopteris flora once formed did not

appear to vary, we have only to explain how it got, one time, into such

remote regions as India, Africa and Brazil, because a continuous ex-

change of the flora would have been unnecessary. Therefore an acci-

dental marine drift of the seeds and the wind, if any were sporogenous,

by way of the Antarctic islands are possible means of distribution, but I

believe that the distribution of the ancestral stock was along the Asiatic-

American “backbone” of the earth and a subsequent similar evolution in

similar environments or else orthogenesis of this stock agrees better with

the known data relating to geographical distribution.

The widely accepted view of a continuous Gondwana Land has been

derived from the static viewpoint of living and extinct animal and plant

94 ANNALS NEW YORK ACADEMY OF SCIENCES

geography, but it is no longer tenable. The separate portions of the

Gondwana Land are, however, more interesting now than ever.

The only places for various past connections which are needed and

almost universally accepted are the following:

Southern South America and perhaps the Antarctic islands.

South America and North America by way of Central America and

perhaps the West Indies.

North America and Eurasia by the way of Greenland and the North

Atlantic, Alaska and Siberia.

Southern Asia and Australia by way of the East Indies.

Kurasia and Africa.**

PERMIAN REPTILES

Does the distribution of the Permian reptiles indicate the existence of

a connection between Africa and South America?

Only a few specimens of Permian reptiles have been found in South

America. Mesosaurus brasiliensis is the best known species. It was

described by McGregor (1909) from the bituminous shales of Iraty,

Parana, Brazil. Stereoslernum tumidum Cope is a closely related form.

It was found in Sao Paulo and comes from the surface of a thin layer of

limestone. Many fragments of it were seen by the writer near Piraci-

caba at a limestone quarry on the property of the Agricultural School.

A few well-characterized marine fossils have occasionally been found in

the series of beds in which Stereosternum is found. 'T'wo more species of

Mesosaurus are known from the Dwyka beds of South Africa.

Mesosaurus is not a diapsid. Its unique vertebre and ribs, as well as

the absence of scales, webbed feet, dorsal but no lateral temporal fenestra,

slender teeth, long snout, etc., separate the genus from all known reptiles.

Von Huene (1910) derives it from some unknown Carboniferous coty-

losaurian. So far, not even the antecedent type, which gave rise on one

hand to Mesosaurus and on the other to Stereosternum, is known. It is

“1 Tt must be granted that of all the evidence in favor of a continuous Gondwana Land,

its flora appears to be the best. But in view of the fact that when it was once formed it

did not appear to change, we may suggest as a future working basis that this flora offers

a special type of orthogenetic development which has been produced from the cosmo-

politan older flora by definitely directed changes in the environment during the formation

of the highlands where it is found. In Australia, this flora appears to have been the

maker of coal during the Permo-Carboniferous. It is also said to be associated with

marine drift and glacial deposits. Hence it appears to be a swamp flora in Australia,

but the presence of thick beds of coal and glacial drift in the same regions does not

appear to harmonize. If this is true, then the Gondwana environments of Brazil and

Australia are distinctly different. A continuous Permian and early mesozoic Gondwana

land is needed no more than a Tertiary or a recent one. We now know, however, that

no Tertiary Gondwana is required to explain the distribution of animals.

HASHMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 95

highly probable that this antecedent type, and not Mesosaurus, was the

form which was distributed.

The exponents of the Gondwana theory must assume that Mesosaurus

originated and died off on the Gondwana continent where its ancestors

are not known, and that it necessitated a continuous Gondwana Land

when its distribution is used to support a connection between Africa and

South America. They must also assume that Mesosaurus was the form

which was distributed and that Stereosternum evolved from it after it

arrived in South America. Two objections can be raised against this

view which are based on positive evidence. First, the nearest ancestral

cotylosaurians known are from the northern hemisphere, and none are

known from South America. Hence the positive evidence in one case

becomes negative evidence in the other. In other words, the positive

evidence is in favor of a northern origin of the ancestral form which gave

rise to Mesosaurus and Stereosternum. Secondly, Mesosaurus was a

good aquatic reptile and did not need a continuous Gondwana Land in

order to get into Africa and South America.

It is evident, then, that the point of origin of the Mesosauria is un-

known. We only know the point of extinction of a few individuals of

three species of the genus. The genus was already very distinct from

other reptiles in the lower Permian, and its ancestral stock could easily

have arisen in the northern hemisphere from some unknown cotylo-

saurian. In fact, Moodie’s paper on the Carboniferous air-breathing

vertebrates of the United States National Museum indicates that this is

probable, because Isodectes punctulatus Cope from the Allegheny series

and Sauravus costei Thevenin from the Carboniferous of France very

remotely point back to Microsauria on one hand and to Mesosauria on

the other. At any rate, the Mesosauria must have originated before the

beginning of the Permian, and the point of origin could have been in the

northern hemisphere just as well as in the Gondwana continent. The

fact that the Mesosauria are known only from Africa and South America

is in favor of the latter view, while the nearest related antecedent types

of Carboniferous Cotylosauria are in favor of the former.

Even if we assume that the point of origin of the Mesosauria was in

either Africa or South America, their distribution offers absolutely no

argument for a connection between these continents, because the Meso-

sauria were aquatic reptiles, as is shown by their long snout, long needle-

like teeth, lack of scales, dorsal position of nares, unique ribs and webbed,

paddle-like feet. McGregor’s reconstruction of M. brasiliensis indicates

that it could not have traveled overland, and inasmuch as it appears to

me that no river could have flowed from Africa into South America, or

96 ANNALS NEW YORK ACADEMY OF SCIENCES

vice versa, during any past epoch, Mesosaurus would have had to enter

the sea in order to get into both Africa and South America.

On account of the intercalated marine Permian in the region where

Mesosaurus is found as well as the underlying limestone containing

marine lamellibranchs where Stereosternum is found, I am inclined to

believe that Mesosawrus was at least semi-marine, if not entirely marine.

Its needle-like teeth strengthen this view, because they are adapted to

eating soft animals, which must have been far more abundant on the

surface of the sea than in the shallow Permian swamps of Brazil, which

became dried up again and again, at which time Mesosaurus would have

been pushed down to the coast. There is, then, little or no doubt that

Mesosaurus could live both in salt and fresh water just as Manatus, and

originally Jnia, which are now found in Rio Amazonas. This being the

case, it could easily have extended its range across the Atlantic, because

it was a good swimmer. It could also have gone by way of the European-

American coast or from the nearest Antarctic islands to southern Africa,

where the distance would not have been much greater than traversed by

the giant tortoises (Testudo) or the semi-marine Jguamde (Ambly-

rhynchus cristatus) of the Galapagos Islands.*?

The mere fact that only Mesosaurus, the best aquatic form, out of

sixty-nine genera of Permian and early Triassic reptiles recently enumer-

ated by Broom for South Africa, has been found in South America, is

strong evidence that no connection existed between these continents.

The absence of this vast array of land reptiles from the corresponding

Permian and Triassic deposits of South America is negative evidence,

but it appears to me to outweigh the positive evidence of the marine or

semi-marine Mesosaurus.**

Scaphonyz fisher. Woodward from the Triassic of Rio Grande do Sul,

Brazil, is another form which has been used to support the idea of a

connection between South America and Africa. According to von

Huene, the known fragmentary data, which have been derived from verte-

bre and foot bones, indicate that Scaphonyx is distinct from Hrythro-

suchus of South Africa. He also thinks that both Scaphonyx and

Erythrosuchus are related to several forms found in the Triassic of North

America. A form of Hrythrosuchus is also known from Europe. Here

again the ancestral stock, which was widely distributed and gave rise to

these genera, is not known. Hence the fact that Scaphonyx of Brazil

has a related genus in Africa is not evidence that these continents were

connected, because it also had related forms in Europe. The great ab-

#T do not believe that the Galapagos Islands were ever connected with South America.

4 It is to be noted that the region of the Permian deposits of southern Brazil has been

inhabited longer than southern Africa, and it is nearly always the natives and not the

few scientific explorers who first find strange animals and fossils.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 9%

sence of Permian and Triassic South African land reptiles from South

America indicates that Scaphonyx evolved from some northern ancestor

from which Erythrosuchus also descended.

Broom and others have attempted to show that the Permian reptiles

of South Africa and Texas are related. There appears to be no doubt

that these reptiles are related, but there is still very diverse opinion as to

how nearly they are related. In the conclusion of this interesting paper,

Broom suggests a possible scheme of distribution of the Permian reptiles.

He thinks that it is probable that a Lower Carboniferous land vertebrate

fauna existed in northern South America. This primitive vertebrate

fauna included among other forms temnospondylous amphibians, primi-

tive cotylosaurians and primitive ancestral pelycosaurs. He presumes

that this fauna ultimately migrated on one hand to North America and

on the other hand over the Gondwana Land to Africa. In order to

obviate the absence of all early Permian reptiles in South Africa except-

ing Mesosaurus, he further presumes that Permian glaciers of Brazil

prevented the other primitive reptiles from reaching Africa until middle

and late Permian.

The above view is possible, but not probable, because it is based on

the following assumptions: (@) a continuous Gondwana Land; (0b) the

existence of lower to middle Permian glaciers in the region of the alleged

trans-Atlantic Gondwana, and not in region of northern South America,

which was with all probability the highest point at that epoch, and (c)

the existence of primitive Carboniferous reptiles and temnospondylous

amphibians which are not known from South America. We conclude,

therefore, that there is little or no evidence in favor of the exchange of

Permian reptiles between South America and Africa by way of a con-

tinuous Gondwana Land.

In view of all this, the only suggestion which appears to agree with

the known facts of geology, paleontology and the Permian environmental

complexes is that the primitive Carboniferous reptiles, from which the

Permian fauna evolved, originated in the northern hemisphere and were

pushed south from Eurasia into Africa, where the descendants retained

certain primitive characters and evolved along similar lines in such a

way that they more or less remotely resemble the descendants from the

same primitive stock which lived in Texas. Mesosaurus is an aquatic

and at least semi-marine form, and does not lend any positive support

to a Permian connection between Africa and South America,** because

44The distribution of the extinct and living side-necked turtles (Pleurodira) offer

another case of the same principle. The pleurodirans are now found only in the southern

hemisphere, but they were very abundant in the Cretaceous of the northern hemisphere,

where they probably already existed in the Jurassic. The two other groups of “shelled

turtles’ (Cryptodira and Trionychoidea) also fit into the scheme of a northern origin

and distribution of land animals.

98 ANNALS NEW YORK ACADEMY OF SCIENCES

none of the typical South African Permian land reptiles have been found

in South America.

MAMMALS

Is an Antarctic connection between Patagonia and the Australian

realm needed to explain the distribution of any of the South American

extinct mammals? :

The best evidence which has been used to support the Antarctica

theory is derived from the mammals. It is the best evidence, because

slow-moving mammals need land connections more than do either flying

or aquatic animals and because the Tertiary record of the mammals is

fairly well known. There is, however, a great blank in the fossil record

in the entire lack of pre-Oligocene mammals of Asia and northern South

America.

The absence of pre-Oligocene animals in both Asia and northern South

America is either due to imperfection of the fossil record or to the lack

of exploration, because the existence of pre-Oligocene mammals in North

America, Patagonia and Africa could not be explained unless the mam-

mals entered both Asia and northern South America; for otherwise we

must assume the separate origin of mammals in two or three different

places. The works of Sclater, Wallace, Lydekker, Matthew, Osborn and

others indicate that the most of the orders of mammals directly or indi-

rectly originated in the northern hemisphere, which has embraced the

bulk of the land at least during the age of mammals. It is true that

South America and Africa have been separate centers of origin of many ~

mammals, but even many of these can be remotely traced back to the

northern hemisphere. The presence of primitive mammals in the Tri-

assic of North America and the Jurassic of North America and Europe

taken in connection with the geology of Europe is sufficient evidence to

show that the pre-Oligocene animals must have existed in both Asia and

northern South America. The distribution of mammals, as I see it, in-

volves, unfortunately, exactly the above regions from which we have no

fossil evidence. These are transitional regions between the northern and

southern hemispheres. Until the known fossil-bearing region of Bahia,

Brazil, is examined and until mammalian fossils have been found in the

early Eocene of northern South America and southern Asia, the distribu-

tion of the Mammalia will never be satisfactorily settled. Nevertheless,

on account of its profound geological significance, I think that a brief

re-examination of the materials of the distribution of South American

mammals should be attempted.

The only support for the Antarctica theory from the standpoint of the

Mammalia is derived from the affinities in the common presence of both

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 99

polyprotodont or carnivorous forms, allied to the existing Tasmanian

wolf (Thylacynus) and of the small diprodont herbivorous forms (Ceno-

lestes) very remotely allied to the phalangers and other Australian

diprotodonts.

The researches of Broom, Gregory, Dieder and others seem to favor

the view that Cenolestes is an independent offshoot of the polyprotodont

type which was present in the Eocene of North America and the Oligo-

eene of North America and Europe. It is also to be noted that the

Patagonian cenolestoids (Hpanorthus and its allies) show no clear evi-

dence of close relationship with Australian diprotodonts.

There can be no doubt that the sparassodonts are true polyprotodont

marsupials, as shown by Sinclair. They also agree with the Tasmanian

Thylacynus in certain characters which have been assumed to indicate

that they belong to the same family. Dr. Matthew, however, is now of

the opinion that these few characters have probably arisen independently

in the Patagonian and Tasmanian genera by virtue of parallel evolution

from primitive didelphids of northern origin.

The Tasmanian and Patagonian genera are the end result of cenotelic

evolution. It was not these genera which were widely distributed, be-

cause there are none in common. It was their ancestral stock, if they

are genetically related, which became widely distributed.

In the case of the cichlid fishes, I have shown that it was not any of

the living genera which were distributed into Africa and South America,

but it was a primitive form. Could not this ancestral marsupial, from

which Thylacynus on one hand and the sparassodonts (Borhyena, etc.)

on the other evolved, have originated in the northern hemisphere from

some primitive northern polyprotodont during the Mesozoic to -early

Hocene? At any rate, it is not yet known from either Patagonia or

Tasmania.

This primitive ancestor could have been pushed out of Asia into Aus-

traha and out of North America during the late Cretaceous to early

Eocene into South America. Then similar evolution in similar environ-

ments would easily account for the rest of the puaulasity of the Pata-

gonian and Tasmanian Thylacynide.

Until it has been definitely shown what this primitive ancestor of the

Thylacynide was and where it originated, it appears to be useless to

reconstruct the surface of the earth from such evidence.

It is interesting to note that the evolution of the South American

mammals agrees in a general way with Schuchert’s view of the connec-

tions which have existed between North and South America. The first

connection existed from the late Cretaceous to the early Hocene, and then

100 ANNALS NEW YORK ACADEMY OF SCIENCES

a separation ensued until the Miocene, after which there has been a per-

manent connection. It is exactly from the Eocene till the Miocene that

South America evolved its typical mammalian fauna, whose last remains

are the anteaters, armadillos, cenolests, sloths and a few tropical mar-

supials. ‘This indicates that the primitive ancestors of these animals

along with others entered South America during the Cretaceous to the

early Eocene. It was during late Miocene time that the second important

change in South American mammalian life took place. This invasion

was without doubt from the north. The third wave was also from the

north. It was composed of man and his domesticated animals. The

replacement of the older fauna by the later invasions is still seen on all

hands in different tropical animals, which still retain the old paleotelic

northern characters which are, however, more or less masked by the

specialized cenotelic characters.*®

In view of all the preceding, the writer, while still in the field, changed

his previous views concerning all of the hypothetical connections between

South America and the eastern hemisphere, and he now believes that all

of the South American animals originally came from North American

stock.

IT am also inclined to believe that the evolution of paleotelic characters,

especially of families and orders, has taken place faster in the northern

than in the southern hemisphere. This is indicated by the fact that

many tropical animals are often a few geological ages behind their north-

ern living or extinct relatives. The edentates, monotremes, ratite birds,

many South American birds (screamers, seriamos, sun bittern, ete.), the

characins, dipnoi, crossopterygians and osteoglossids (fishes), South Afri-

can secretary bird, note Aardvark (Orycteropus), scaly anteaters (Manis),

tapirs, camels and many marsupials are examples of tropical animals

which are a few geological ages behind time. This retarded evolution of

paleotelic characters in the southern hemisphere may be due to a greater

stability of the vast Plano Alto of South America.

It is not, in my opinion, the stable portions of the earth which have

produced the bulk of evolution, but it is the ever-changing regions either

by elevation and submergence or tremendous changes produced by ero-

sion, like the recent formation of the Amazon Valley. These violent

changes produced in the environmental complexes appear to pull the

trigger of evolution. Inasmuch as geology shows that more radical

45In taking this view, I have assumed that Matthew and Gaudry are correct in con-

sidering the Patagonian beds to be later than the upper Cretaceous. Roth, however,

considers the Notostylops beds to be upper Cretaceous, and Ameginho considers them to

be still earlier.

HASEMAN, GEOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 101

changes in the environments, including the climate, have occurred in

North America than South America, it appears probable that paleotelic

characters evolve faster. This belief may seem absurd, yet if it is only

in part true, it has a profound significance in correlation, and especially

in determining the exact age of the fossil beds in South America.

If it is true, then older-appearing South American beds are in reality

much more recent. At any rate, this impression demands careful study,

and especially in the case of the invertebrates.

I believe that the North Polar theory of the origin of land animals

expounded by Haacke and in a general way supported by Wortman,

Sharff and more recently by Matthew, is the view which agrees best

with all of the known facts of geology, paleontology and zoology. Ruti-

meyer, Huxley, von Ihering, Forbes, Ortmann, Hedley, Sinclair, Ame-

ginho, Osborn and others have maintained connections between Australia

and Patagonia, but their evidence has been derived almost entirely from

the static viewpoint of zodgeography, which, as Tower has well said, is a

dead and profitless pursuit. Besides, there never has been a general

agreement between any of these authors either in regard to exact position

or time of existence of the connection. They have also utterly failed to

show how and why just certain animals were able to get across the con-

nection. Why, for example, did not edentates and other early Tertiary

mammals of Patagonia also get into Australia? Would not such a con-

nection have had a barrier? - Besides, the distance across this south polar

continent is not small. They also do not attach much importance to the

strong geological evidence against such connections.

The ideal northern marsupial from which we could easily derive both

Thylacynus and the sparassodonts is not definitely known to exist, but it

is also not known in either Patagonia or Tasmania. In fact, I should

expect to find it in Asia and northern South America, both of which

places are entirely unknown from the standpoint of primitive mam-

malian paleontology; but even if the necessary ancestor is never found,

it will not be the only gap left open in paleontology.

The indecisive evidence used in support of the Antarctica theory does

not appear to me to outweigh the fact that neither the deep-sea sound-

ings, the trend lines, the lack of islands, location of Archean rock nor

the location of known marine formations even vaguely suggest a Pata-

gonia-Australian connection. Besides, such indecisive biological data are

not as weighty as the vast array of data in favor of the persistence of the

continents and the great ocean beds, so ably defended by Sir John Mur-

ray and others. The deposits in the great ocean depths like those be-

tween South America plus the Antarctic islands, Africa and the Aus-

102 | ANNALS NEW YORK ACADEMY OF SCIENCES

tralian realm have never been found in the whole geological series of the

continental shelves. Also, the great number of shark teeth found in

abysmal depths indicate vast time for deposition. Until all such data

and the theory of isostacy have been satisfactorily accounted for, it ap-

pears to be useless to continue hypothesizing land-bridges. At the present

stage of our knowledge, we do not need an Antarctic land-bridge, but we

do need both dynamical data and more careful field work in northern

South America and southern Asia before we can definitely settle the

distribution of mammals.**

SUMMARY

We have seen from the location of the Archean and early Paleozoic

rocks that about the present outline of South America has always existed

and that the lines of weakness and strength in its crust are usually par-

allel to the coasts. Hence, the invasions of the seas have, in most cases,

been in a general southern-northern direction and not east-west. The

location of the deposits left by the invasions of the sea has forced us to

deny the existence of Archiguiana, Archamazonia and Archiplata as

maintained by some of the exponents of the Archhelenis theory.

The outlines of the Plano Alto, which was deposited in a continental

Permian inland basin, has been given, and the general dip of its surface,

its lack of past Paleozoic marine deposits, location of surrounding

Archean mountains and marine sediments and the Tertiary rise of the

Andes indicate the reversal of the Amazon during the later half of the

Tertiary epoch. The eastward movement of the mouth of Rio Negro and

the single channel of the Amazon in region of Obidos, where remains of

the Plano Alto approach the river, indicate that this is near the old

divide which has been washed away. The unique marine or brackish

water fossils of Alto Rio Amazonas apparently lived in an arm of the

ocean (Hast Andean Sea), which probably extended south, lost its con-

nection and finally disappeared with the Tertiary rise of the Andes. It

was also suggested from the character of the overlap that no great exten-

sion of land to the east of the present coast was needed to form the sedi-

ments of the Plano Alto and that great altitudes probably existed in

eastern Brazil and Guiana during late Paleozoic times.

The southeastern Brazilian coast appears to be very old and remark-

ably stable. It apparently never extended more than about 100 miles to

the east of its present location. The fringe of upper Cretaceous deposits

46 The evidence from the standpoint of the Mammalia for a Tertiary Archhelenis, 7. e.,

a connection between South.America and Africa, is given in the “Age of Mammals,”

which shows that such a connection did not exist.

HASEMAN, GHOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 103

along its northern portion, the fan-shaped trend lines west of Pernam-

buco which fade away toward the coast and are roughly comparable to a

somewhat analogous condition in the region of La Paz, Bolivia, its recent

slight elevation, its lack of numerous islands separated by comparatively

shallow sea and its abruptness being primarily due to erosion of Paleozoic

and post-Paleozoic land deposits and not to post-Paleozoic faulting, indi-

eate that South America was never connected with the eastern hemi-

sphere. This view is further strengthened by the absence of deep-sea

ooze, etc., from the continental shelves and the abundance of sharks’

teeth in the great ocean depths which indicate a vast antiquity for the

abysmal depths.

We have noted many facts concerning the South American topography,

the most interesting of which were the cases of stream piracy existing

between Rio Orinoco and Rio Negro, and Rio Sao Francisco and Rio

Tocantins. The Paraguay River is not connected with the Amazon.

These facts, taken in connection with waterfalls, altitude, swamp produc-

tion, erosion and the composition of environmental complexes, have led to

some interesting results concerning the distribution of the South Amer-

ican fishes.

Using the cichlid fishes only as the best known family of South Amer-

ican animals, it has been shown that isolation or barriers and interming-

ling or river connections utterly fail to explain their distribution. It was

found that the present distribution of the fishes is correctly explained by

the organic complex of the more generalized highland genera (which are

small in size and naturally widely distributed because the Plano Alto was

formerly a continuous unit) and by the action of the environmental com-

plexes on this stock. In other words, when the common ancestral forms

arrived in similar environments, 1. ¢., similar environments were eroded

in the Plano Alto, they evolved along similar and identical lines and in

different environments along different lines.

When we attempted to determine the point of origin and lines of dis-

persal of families and orders, it was found to be absolutely necessary to

invoke the aid of fossils. In doing this, it was found necessary to use

more than single physiological characters and draw a sharp distinction

between paleotelic and cenotelic characters. When this was done, the

fishes evidently point to a northern origin and not to an African-South

American Gondwana origin.

When similar methods were applied to the Permian reptiles, Gondwana

flora, mammals and other alleged evidence in favor of connections be-

tween South America and the eastern hemisphere, the evidence was not

found convincing for a single case. Thus the Permian reptiles, if crit-

104 ANNALS NEW YORK ACADEMY OF SCIENCES

ically studied, offer evidence against instead of for a continuous Gond-

wana Land. In fact, all of the alleged evidence has been derived from

the static viewpoint of plant and animal geography which has led to

many erroneous views of correlation and geology of South America.

There was not found a single case in the evidence for a continuous Gond-

wana of any age or location in which the distributed ancestral form was

actually known.

In view of all this, I have been forced to change my former belief in a

connection between South America and the eastern hemisphere, because

the geological evidence overwhelms the biological hypotheses. The frag-

mentary positive evidence in a few individual cases may not always indi-

cate that this view is true, but when both positive and negative evidence

derived from botany, zoology, paleontology and geology is carefully

weighed and due allowance is made for the imperfection of exploration

and the imperfection of the fossil record, the evidence is decidedly against

the hypothetical connections.

I therefore believe that continental forms have originated and dis-

persed over three great tongues of land which have always extended south

from the northern hemisphere. These three great tongues of land have

been connected and disconnected from time to time, and it is possible

that they were connected at the south pole at some time previous to the |

Carboniferous epoch, but so far there is little or no evidence for such a

view.

There have been from time to time possibilities of plants and animals

interchanging between these three tongues of land by way of the north-

ern hemisphere.

In a word, we do not accept the theses of hypothetical land-bridges

and invasions of the sea; we fail to appreciate the weight of the evidence

in favor of these theses, and we look forward with keen interest to the

results of coming years in field work and in dynamical studies especially

in the regions of zoddistributional transitions, 7. e., Central Asia and

northern South America.*?

‘7 Tt is in a way extremely unfortunate that so much work is done in regions like Pata-

gonia and southwest United States in order to prove a theory which can only be proven

by hunting the deposits in northern South America.

HASEMAN, GHOGRAPHICAL DISTRIBUTION IN SOUTH AMERICA 105

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110 ANNALS NEW YORK ACADEMY OF SCIENCES

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112 ANNALS NEW YORK ACADEMY OF SCIENCES

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ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXII, pp. 113-133, Pll. XVIEXIX

Editor, EpMuND Otis Hovey

SEX-LINKED INHERITANCE IN POULTRY

T. H. Morcan anno H. D. GooDALE

NEW YORK

PUBLISHED BY THE ACADEMY

23 JULY, 1912

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Narurat History, 1817-1876)

OFFICERS, 1912

President—Emerrson McMitxin, 40 Wall Street

Vice-Presidents—J. EDMUND WoopMAN, FRrepERIc A. Lucas,

CHARLES LANE Poor, R. 8. WoopwortH

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SECTION OF GEOLOGY AND MINERALOGY

Chairman—J. E. Woopman, N. Y. University

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[ANNALS N. Y. Acap. Scr., Vol. XXII, pp. 118-133, pll. NVII-NINX, 23 July, 1912]

SEX-LINKED INHERITANCE IN POULTRY

By T. H.. Moregsan anp H. D. GooDALE

(Presented before the Academy, 8 April, 1912)

CONTENTS

Page

SUE ste ( UA Ls1 GENE eee ea near eP ei tacia! Sle) ceo ersten ccve Shove) Peiieca)ven've: 3 eis! en's: o'9:l6 ie) 6) ep! ea 113

Crosses between Plymouth Rocks and Langshans.....,.................-. 114

IDSSCENTMNOINIOE TKS KAN Sooecogacnooeeooe den so dddoooon Tage eet aod ener ak cual ood 11+

EMIS tO IVAO tab MCT CO US ayerdisrar terse syaieier clatlntthcle io. eke le eeayne Sustcesie is a!e ce Seisvey eves 115

Source of breeding stock used.................6., Make Nasheed ea eee ctaice os 115

PUT eaten seater ec te apn = ey atrag ses ac Nicene Sie ube ehe erie Srai'a neers Ga islole! eheveyale es 115

Explanation of the symbols used in interpreting the results.......... 116

Parental matings...........-. cee eee eee eee eee eee eee cece eee 118

1B, TURGUENTEIS. Sb tiesnicios O's See Cae EDO OREO OD a een ncaa er 118

IBAGIR TWNIMNES C56 das Bid an Cle.c BiG oid COC OG Ota DI IO pIOI it ai eno Io ini Carre 120

SITUOMUMMATAY ond SSE GSCI eek ERAS IS OP CR EERE Ree rear Beal et a cater kis 121

Meseripmoncvok By Adult Rlumace.-. o2.5. 2c .ccnssseessdacnes sevens 12.

SS eaira Kee (20) Oder epenenecteate oicictarersyaicke cushanciats lie close cro oe skin aid aus'b.@ ww sidie Se Siar 123:

ESCO CH re tepeterecet st cert cha arate ay ereiare suse wise sae claves a aisrdnay so ale angome eed alee au ese 1256

Down colors............. > GeO GiCh OC SmNCEO ES CC CRERERACRS CMTe eR ae ee ES

Crosses between American Dominique females and Langshan males...... 128°

SAE Ti ee CMLETAGI Ol oes crores eperatevatece? acl ers (esere cis Weave elelsvegiaicia vce sie abaule dle ses 128

F, generation from Langshan ¢ by Dominique 9.,.................. 128

F, generation eee R een ace ICIN SEMEN rarer ae aiavare laielars vie accuse a aieisis vais se waa 128

Back cross of F, ¢ (barred) to Dominique 9...................... 128

Back cross of Hi, black 9 to Langsham @.......-........c.2-..00000% 129

Oper teatures’ Of the CrOSSESi oo. obec de ce yee eee eee se cew esses 129

WIKeR heaters IMy WINGS tine cs acc eas ees ccs case sve dens ee mele eet ewses 130

COOP OF WEBI SS Ss Sola Gb EG RO SEES Oe Sra sa ee 131

Color OE TOM Ses Sere Go is. a Secs ecel OEN OREM Sep ence ene er =e ee ort 131

MEO GalerCONSIGCTALIONS. cc. 626s As eevee cc sess scenes ce decsaeceeewsec 131

SSH fom CHOI eed Ju NVM eae em at oe ai larie> hela) a Aiicto ove, bio Gia ge wiaete gh mieinis Biase ee 133

INTRODUCTION

In 1908, W. J. Spillman pointed out that, according to a breeder,

when Plymouth Rock females are bred to Langshan males all the females

are black and all the males are barred. As far as the evidence went, it

seemed to show, as he pointed out, that the case was comparable to that

of the moth, Abraxas, described by Doncaster and Raynor, and of certain

crosses among canaries described by Miss Durham.

(113)

114 ANNALS NEW YORK ACADEMY OF SCIENCES

With the intention of examining further the report cited by Spillman,

and of testing, by further combinations, the offspring of the first genera-

tion, we began the following experiments in 1909, using Barred Ply-

mouth Rock and Langshan fowls. We undertook also to extend the

experiment by using another breed of barred poultry, the American

Dominiques.: It is currently stated that Dominiques (but not American

Dominiques) occur in the ancestry of Plymouth Rocks. We wished to

see whether “sex-limited” or “sex-linked” inheritance is found also in

this other race. Plymouth Rock-Langshan crosses have been made by

one of us (Goodale) on the experimental farm of Mr. B. B. Horton, to

whom we are under many obligations for opportunities to carry on the

work. The Dominique-Langshan crosses were made by the other (Mor-

gan) at Woods Hole during the summers of 1910-11. In the meanwhile,

Pearl and Surface (1910) have described the results of a cross (and its

reciprocal) between Barred Rock and Cornish Game. Goodale (1909,

1910) has given briefly some of the results obtained when Barred Rocks

are mated (reciprocally) with Buff Rocks and when Brown Leghorns

are mated with White Rocks. Hadley (1910) has called attention to

similar results published by Cushman in 1893. Davenport (1906, 1909)

has described various crosses to one of which certainly (White Cochin by

Tosa) and to the others less clearly may be given the same interpretation

that apples to the results described in the other papers mentioned above.

These crosses all involve the barring factor. Sex-lnked inheritance of

other factors in poultry has been noted, not only by several of the above

writers, but also by Hagedoorn (1909), Sturtevant (1911) and Bateson

and Punnett (1908). To Bateson and Punnett is due the explanation —

of the phenomena of sex-linked inheritance for poultry. More recently

(1911) these authors have published a complete account of the inherit-

ance of a factor derived from Brown Leghorns which affects the patency

of the type of pigmentation characteristic of the Silky fowl.

CROSSES BETWEEN PLymMoutTH Rocks anpd LANGSHANS

Description of the Breeds.—F¥or a detailed description of the breeds

under consideration, reference must be made to the various standard

works on poultry. In this paper, only a very brief statement of the

chief characteristics involved in the cross will be given.

The Black Langshans (Plate XVII, figs. 2 and 3, and Plate XVIII,

fig. 1) are uniformly black, varying somewhat in brillianey in different

regions of the body. The shanks, too, are dull black; the bottoms of the

4 The American Dominique is a younger breed than the Barred Plymouth Rock.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 115

feet are gray. The shanks, moreover, are provided with several rows of

feathers, or boot, along the outer edge. The comb is single.

The Barred Plymouth Rocks (Plate XVII, figs. 1 and 4; Plate XVIII,

figs. 3, 4, 5, 6) are black and white,” the two colors being arranged in

alternate bars across the long axis of each feather. The bars vary some-

what in evenness, width and depth of color from individual to individual,

and also in different sections of the same bird. Although the American

“Standard of Perfection” requires that the two sexes shall be alike in

color, the males vary from a darker to a very light color, often appearing

very light gray, while the females, though to a less marked extent, vary

toward a darker shade. In other words, the breed tends strongly toward

a sexual dimorphism of color, with indications of a secondary dimor-

phism within each sex. The comb is single, and the yellow shanks are

free from feathers.

History of the Breeds —The modern Langshans are the direct de-

scendants of a very old race brought from the interior of China. The

Rocks, on the other hand, resulted from a mixture of several races of

fowls about forty years ago, which have been gradually brought to a

high degree of perfection. The history of the barred character with

which we are chiefly concerned is obscure, but evidently it is of very

great antiquity, for barred or “cuckoo” birds are to be found in many

European countries, Russia included. Brown (1906) states that the

plumage of the “Siberian Featherfooted fowl is generally white, whilst

others have cuckoo plumage.” He notes also that this variety is said to

be of ancient lineage. Wright (1902) states that it is probable that the

“original Chittagongs, or at least their crossed offspring, were of an

‘owl’ color as described, probably what we now know as cuckoo or barred.”

The Chittagongs came from the district of that name in the upper Malay

peninsula. An exhaustive search would probably show that barred fowls

have been recorded from southeastern Asia.

Source of Breeding Stock Used—The Langshans came from P. P.

Ives of Guilford, Conn. ‘lwo of the three Barred Rock males and one of

the females were of the well-known Latham strain, but obtained from

R. C. Goodale. Four of the barred females were the progeny of the

Latham hen by a White Rock male, one was an F, from a similar mating,

and one was a pure bred female from a Stamford breeder. The White

Rock male is known to differ from the barred birds chiefly in the absence

of the chromogen factor.

Matings—In the majority of these matings, the progenies of the indi-

vidual mothers have not been kept separate. The determinations of the

2 Fanciers prefer to speak of both these colors as grayish.

116 ANNALS NEW YORK ACADEMY OF SCIENCES

presence or absence of barring, unless otherwise stated, was made on

newly hatched chicks, or those of full term which failed to hatch. This

method of determination is made possible by the presence of a gray

occipital spot on those chicks which will become barred adults. A full

discussion of the point, however, will be given elsewhere. Inroads of

vermin, largely rats, have limited the number of which the sex was de-

termined. A description of the adult hybrids is deferred until after all

the matings have been described.

Explanation of the Symbols used in Interpr eting the Results.—It was

pointed out by Spillman, following Bateson, that sex-linked inheritance

in poultry could be accounted for on the assumption that the female is

heterozygous for sex and the male homozygous, and that when in the

female, the barred factor alone is present, it is repulsed by femaleness.

We may give this interpretation a more concrete form, if we assume that

the factor in question is not carried by the same chromosome that carries

the factor for the female sex; 7. ¢., in the heterozygous female the chro-

mosone that carries femaleness also lacks the factor for barring, and its

mate that lacks the factor for femaleness carries the factor for barring.

No interchange between the chromosomes (if two really exist) can take

place, perhaps because they fail to pass through those stages in synezesis

when such a process becomes possible.

If F = female, f its absence or male; B = barred, b its absence ; N=

black, then the formulas for the barred ok will be:

asi CRO cwiarever st suet svayatoesaiatitca cs iskehartie 4 Se muewere FENb {NB

: Omaha oeds Sie isast aie «Sie aes eA {NB {NB

Bacio POC nears eee: eee dmmne ra CELLS 3 FNb fNb

CUE ey tl ak es we ae) ee fNb {Nb

Whether the female-producing gamete of the Barred Rock really car-

ries black or only the absence of barring will not affect the nominal

results here recorded, but other experiments to be described by Goodale

will show that “black” is probably present.

In order to see how these formulas apply to the crosses under con-

sideration, let us take first the case of the cross between the Langshan

hen and the Plymouth Rock cock (fig. 1). The formule are as follows:

amg shan: O03 ae eee orto recess cui sais FNb fNb

Barred ‘Rock «8.0. 5 eee Se eee {NB {NB

a Lone Qs ls Goer ee DS 85 FNb fNB_

Ee Sen ee fNb fNB

MORGAN AND GOODALE, INHERITANCE IN POULTRY 117

IBIENOG ER: O.'5 hg bbe ded GOR Se eC OER Rene eee FNb fNb

USD ES crates Ommeveyenee i crstijehe el Sic. waitose: ciclg esto silts se Seis ENb fNB

2 i; MEST Oe Uo Net spel eiaray sun wie eral BALS {NB fNB

aren mieten reese echey se Ss0s) sau cajie es Sos esas: wale {NB fNb

Barring is dominant to self color, as is shown in the last case, where in

the F, generation all the offspring are barred. In the second generation,

Fig. 1.—Cross of Plymouth Rock ¢ to Langshan 9

there oceur barred ¢ and both barred and black @. The grandmaternal

color, black, appears in the grand-daughters and not in the grandsons.

The reciprocal cross between the Barred Rock female and the Lang-

shan male (fig. 2) may be represented as follows:

Ang C EPENO CRY OMe cose eo): ajlatetste eheus sieht oie: setae FNb {NB

HATS IN TTD Meme A Reyer ey ecroncate, sais tel oy sirshoueie io) spt crap-se tiene fous fNb {Nb

eae ee ee aia. marie ..... FNb £Nb

pee PE MUTT CLC Ret shes eos ni Gil ih cna fa Mn wie one edo eneeelelonene {NB fNb

( SATCU See O rite so) cen stso tes Seicususerace Sie) sus. atvcene maerets ENb fNB

I BSI @ Ee Saloval a raat ieseerie soa ack ce a ae ame coe EE VS ones ae FNb fNb

2 4 AEC Me a st aiaie statis Srasasrouslieile edocs ious giao leapsiiomenie os f£Nb {NB

ISIC) Gees econ e cucenaro uo uromapa cd see {Nb f{Nb

118 ANNALS NEW YORK ACADEMY OF SCIENCES

Parental Matings.—(1) From the five Langshan hens by a Barred

Rock male, there were 34 young, all barred: 12 were females and 8 were

males (fig. 1):

FUND? SGN eecse artes aos iS aceievel eae eee Langshan 9

TINUE SIENYS Sepatcrscarcscca sare ene. wc eeoei sy oh oacaiieballe Barred Rock ¢

BUN fo eee DINU Sf aie senstere seacnisne seater ostiane voneccenrete Barred @ 12

NIB NIB CS. o crate eae iawn ei reeeke e a 8

Fic. 2.—Cross of Langshan g to Plymouth Rock e)

(2) From the various barred females bred to a Langshan male, there

were 20 barred (15 4 ) offspring and 25 black (162) (fig. 2):

BN ENB Cee ee eae eee enter eee ae Barred 9

fNb ENDS 52) Wee eee ee Langshan ¢

NENG re INO 3s ee ee ae eet Black 92 16

PINIB GUND i ate ele eee oer ae Barred @ 15.

PF, Matings—(3) Four barred F, females from (1) were bred to a

barred male, No. 568, from (2). This was done because the only adult

MORGAN AND GOODALE, INHERITANCE IN POULTRY 119

male, No. 784, from (1) did not mature until long after his sisters were

laying, while a change of residence on the part of the writer prevented the

accomplishment of the inter se mating. From the cross-mating, however,

there were 25 barred (12 ¢ and 9 @ ) offspring and 13 black (8 ) (fig. 1).

Expectation on the Spillman-Bateson hypothesis is 2814-914. One indi-

vidual, a male, with the gray spot reduced to a few plumules was excluded

from the count as doubtful.

( IEINID TENIB ceeceoance suceccsacgenode Barred 9?

BE OND) cee Shen see tara

IRUNG SNUB 25 28 nate ere ele yorseetenctene sce ,0 iotee Barred @ 9

MMBEBN es CEINIDIR. 0 care tc once eases Black 2 8

Fe fNB

| fNB

Hic. 3.—Cross of barred 4 to Fy black 9

(4a) The 6 black females from (2) were mated to a litter barred

brother, No. 569, giving 41 young; 16 of which were black (86, 62 )

and 25 barred (66,109). (40) Later, they were bred to No. 784,

giving 22 young; 7 were black (06, 192) and 15 barred (56,192).

The combined results of these matings were 63 young, of which 23 were

black (86, 72) and 40 barred (116,112) (fig. 2). The departure

from the expected ratio of 3114 is considerable.

120 ANNALS NEW YORK ACADEMY OF SCIENCES

r } TEND END ke Oe ciones ete eredeters eaisteemn Black 9

©) Os SfINTES A ethIND sis Sree eaten esc Bee sees Barred ¢

( FNbB fNB.........---. ee eee eee eee Barred @ 10+1

RUN DS ao aeNiby ee ss catches Mem ameleee Grae Black 9 6+ 1

Ea AO AEN EN se ae oe Barred @ 645

EN De CEN De cd.c.ce-ae cet hee esate he egere Black ¢ 8+6

Back Matings.—(5) The 6 black F, females used in (3) and (4)

were bred to a pure Barred Rock male and gave 9 barred young, fulfill-

ing expectation (fig. 3).

NID aeBIND octane estes eee eee Black @

ENB) ENB aaccstis cy sitet eieisteraioseooete re Barred ¢

TUNIS US ect os clea eee eetre ee aie Barred 9

PINT: SEIN a ercierend Si chaternsee ekercho cael ceeveaaione Ss a

Vic. 4.—Oross of Langshan @ to Fy, barred 9

(6) The 4 barred F, females used in (3) were bred to a pure Lang-

shan male. Of the 30 offspring, 14 were barred (14,12 ) and 16 black

(06,32). This corresponds closely to the expected equality in ratio

(fig. 4), except for the possible barred female. The determination of

sex im this case was made on rather poorly preserved material.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 121

TN) aiN1B Ges jogo aro coated Gh Oenioro Barred 9

(ENTS. ENID cedecan se coos coped ece pone Black ¢

ify JENID goeeeneougcnococbodep oda Black 9

(NIE ENO? G4 cedure sue eoooos Goon Ot en Barred ¢

(7) Several of the parental barred females were mated with No. 568,

barred F, @, giving 8 barred (32,14) and 3 black (12 ).

EWING Tove ete Nilo oes apie open laut ashy clickel oMelloneers pees Barred 9

NIS. OND coneisouoddqaseouuoupeuabeT aad ones

TEIN Loe wae NUS ibe fe ese ye ses nerceriehsy ef elemece pain sretalecays Barred Q 3

IRIN Dy cpa] Ota e's & cuesmrcrche onc eine Ura irae a Black @ 1

if NG ENG ate cicie sieee ib hsloy a ckcie sue. erevese we Barred ¢

FINGES MOEND Le ene aN eA “4 \ =

(8) Four of the parental Langshan females mated to No. 569, barred

F, male, gave 17 barred (62,26 ) and 13 black (44 ; no record for® ).

TRIN OLSEN OSs one eeeorereicio otectorais cea ey oaetrae Black 9@

fNDES eae TING cacerncncre heroes agate sey cqagscls Sree eetens Barred ¢

FUN unmet NES ener echchore are ieraite essen sachs 3 leha vhs Barred 9 6

FUN De eet skarcaec ocgerays 2 stoke cha dae ete ae Black @ ?

TEN O%m COENY uke Oe sea ere rh eR a Barred ¢ 2

ifSIN JO ate aN Dicre ates iene sie eee ni aie a aes sens Black ¢@ 4

(9) Two of the barred F, females used in (3) were bred to a Rock

male, not, however, of the Latham strain. There were only 5 chicks, all

barred.

TEIN Dean PINE eae aes Pie fers la) > a iets evare aieleta'e Barred 9

ESTEE FENTR OL E IO eee ee a nr

HINDU ENIBe Eee... eat en eee -.. Barred 9

NTE PEES1S So oe “

Summary.—Expectation in all these matings has been closely fulfilled

(with the exception of No. 4 and perhaps No. 6), on the assumption that

the barred female is heterozygous for both barring and sex, both female-

ness and barring being dominants and that the two factors do not occur

in the same gamete. ;

Description of F, Adult Piumage.—The males (Plate XVII, figs. 5

and 7) resemble one another closely and together with the barred F,

females are very suggestive of the Coucou de Malines, a Belgian breed.

The barring of the individual feathers of the males (Plate XVIII, figs.

10, 13) is less sharp and regular than that of the parental Rocks (Plate

XVIII, fig. 8), while the dark bars tend to run together, particularly in

wings and tail, and at the same time, the light bars become more or less

122 ANNALS NEW YORK ACADEMY OF SCIENCES

smoky. The primaries, indeed, can be called barred only by courtesy, for

the light bars are only represented by white splashes along the shaft

(Plate XVIII, fig. 14). This region, however, is one in which the fan-

ciers have found great difficulty in producing even and regular barring.

One barred male is particularly interesting in that a few feathers show

distinctly the Jungle coloration (Plate XVIII, fig. 12) which probably

exists as a cryptomere in the Langshans.

All the males have numerous feathers wholly or partly black (Plate

XIX, figs. p-w) and this is true also for the barred females. ‘The last,

except for the black feathers, are well barred (Plate XVII, fig. 6) and

can scarcely be distinguished from the parental stock. Even the indi-

vidual feathers, except the remiges and retrices in which the bars run

together, conform closely to the pattern shown by many thoroughbred

Barred Rocks.

The color of the F, black females (Plate XVII, fig. 8) is indistin-

guishable from the parental Langshans.

Comparatively few members of the F, generation reached maturity.

The only points of particular interest are the appearance of very light

as well as dark males, of both black and barred males having a few

feathers showing the Jungle fowl coloration and of dark-colored females

with the barring somewhat blurred.

The non-appearance of game (Jungle) colored birds in F, is due pre-

sumably to the fact that black is duplex in both Rocks and Langshans,

and thus the Jungle fowl color is concealed. There are, however, indi-

cations that black may sometimes exist in a simplex condition among

Rocks; so that, if suitable matings were made, the Jungle fowl color

might appear. Unless the occasional feather showing Jungle fowl color

is due to a simplex condition of black, its appearance may mean that

hybridization in some way has upset the usual complete dominance of

duplex black over the Jungle fowl coloration.*

As already stated, Pearl and Surface have published their results in

crossing Plymouth Rocks and Cornish Indian Game. Our results en-

tirely accord with theirs, as far as inheritance of barring is involved.

They classify their birds as barred and non-barred, ignoring intentionally

the differences among the non-barred birds. Our results are simpler, in

so far as all our non-barred birds are black, but the principle involved is

the same in both cases. Pearl and Surface have also made all possible

back crosses between the parents and the F, generation. Our results are

in entire harmony with theirs, but they have the advantage of a larger

number of offspring in their matings.

3 DAVENPORT, 1909, p. 72.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 123

Shank Color.—The color of the shanks of all chicks hatched was re-

corded, but the color often changes as the birds become older, so these

records prove to be of small value. The change in shank color is

particularly characteristic for the class called yellowish or flesh-colored

black. This class may give rise to all three of the adult shank colors.

recognized, black, gray and yellow. Black-shanked chicks seem always

to develop into black-shanked adults, and while yellow-shanked chicks

probably do not produce black-shanked adults, they may give rise to.

either yellow or gray-shanked adults.

The infantile shanks among the F, not only show the expected classes,

but these classes pass by imperceptible grades into one another. Fre-

quently, one part of the shank, particularly the toes, differs from the

remainder ; while, in many cases, the distribution of color forms a mottled

pattern. The distribution of color upon the toes is likewise extremely

variable and often asymmetrical. In almost every case, however, some

part of the toes is flesh or yellow. This variation is due, presumably, to

some extension or restriction factor. Similar variation in the distribu-

tion of color in F, was also recorded.

The shank color of the F, adults falls into two classes, black and gray.

The term gray is used rather loosely to cover a particular though some-

what variable coloration of the shanks. At a distance, the shanks do

indeed appear gray just as a Barred Rock appears gray, and just as the

“eray” of the Rocks resolves into a pattern on closer inspection, so the

gray of the shanks is not a single or uniform color. For convenience of

description, we may say that the ground color is steel gray, variously

mottled with patches of darker gray or of black. Parts of the shank

often have a bluish cast. The posterior side and particularly the bottoms.

of the feet are somewhat flesh colored. Mottling does not as a rule occur

on the bottom of the feet, so that though the term gray is applied to them

in a later paragraph, it is to be understood that they do not have the

same appearance on the shanks proper but rather are a grayish flesh, self

color. The three classes of black, gray and yellow do not grade into each

other.

The six F, black females had black shanks. The three males and the

four barred females had gray shanks. Apparently, we have here a case.

of sex-linked inheritance. This, however, may not be the case but may

be due rather to the black spreading over onto the shanks just as it often

spreads over onto the comb. - In the barred birds, we may suppose that.

the barring factor operates to prevent the spreading of black over the

shanks, just as it also produces the characteristic barring of the feathers

of the boot. Thus, the colors hypostatic to black are revealed. However-

124 ANNALS NEW YORK ACADEMY OF SCIENCES

this may be, in F, the black birds again have black shanks, but the bot-

toms of their feet, which are usually incompletely covered by black, are

either gray or yellow. The allelomorphs involved, then, are gray versus

yellow (or Gray,, Yellow,, X, No gray., Yellow,), the latter being re-

cessive to the former. Moreover, among the barred birds, only gray or

yellow shanks appear, or in other words, gray-shanked birds always have

gray soles, yellow shanked birds yellow soles, but black-shanked birds

may have either gray or yellow soles.

Since, then, the black-shanked condition is due to an extension of the

general black color of the body, we need consider further only the rela-

tion of gray to yellow, the determinations being made, of course, only

on the bottoms of the feet and when the birds were several months old.

In F,, there were only gray or pinkish gray feet, and, therefore, there is

no evidence that gray is sex-linked. Moreover, since no other color than

yellow appeared in F,, yellow is probably common to both Langshan and

Rocks, so that absence of gray in this case means yellow. In F,, not all

the adults were available for study, as the importance of foot color was

not realized until after many of the birds had been disposed of, but in

17 cases, 13 were gray and 4 yellow. The back mating of F, gray male

to P, gray (Langshan) female gave 6 gray. The back mating of P,

yellow (Rock) male to F, gray (black plumage) female gave 6 gray to 2

yellow. ‘These results indicate, then, that gray and yellow feet (or

shanks, leaving out of consideration the supermelanic coat) behave in ~

simple Mendelian fashion.

We have suggested that black individuals have black shanks, because

a restriction factor is absent from these birds, so that the body color

spreads out as a self color over the shanks. Such a “restriction” factor

would be sex-linked. Is it, then, the same as the barring factor? If it

were a separate factor, we should expect that, in F., a certain amount of

segregation would take place. This has not been observed, so that it

seems probable that the black shanks of the black birds are due to the

absence of the barring factor and the mottled shanks to its presence,

unless some “association” exists. Thus, the presence of the barring fae-

tor results in two (perhaps three) distinct somatic conditions, viz.:

barred feathers and mottled shanks, and, as a possible third, the gray

occipital spot of young chicks. In other words, we have two or more

unit characters resulting from the operation of a single factor.

There are some considerations of a practical nature resulting from the

relations between shank color and sole color which should be mentioned.

If the black color covered the entire foot, we should be unable to deter-

mine what color underlay the black, except perhaps by long-continued

MORGAN AND GOODALE, INHERITANCE IN POULTRY 125

breeding tests. Gray would, therefore, appear to be a sex-linked charac-

ter. In F,, however, the results would appear peculiar, for while we

should have the three classes of black, gray and yellow shanks, the black

shanks would always appear associated with black birds, while gray and

yellow shanks would go with barred birds. This conclusion does not

agree with the results expected when two independent sex-linked char-

acters are involved. In F,, the observed results would be very compli-

cated. A discussion of the various possible explanations which might be

devised to meet the situation would hardly be profitable here, but a com-

parison of the results expected when the color of the soles of the feet is

taken into account with those when they are omitted may furnish the

_key to similar cases.

Booting—The Barred Rocks are typically clean shanked, but occa-

sionally a bird is found with a few “stubs.” The boot of the Langshan

corresponds approximately to that shown in many of the older pictures

of Cochins and Brahmas and may perhaps be regarded as the primitive

type from which the modern highly developed boot of Cochins has been

developed.

For the F, generation, booting was recorded on the chicks as “present”

im all cases but two. These two occur among the first four recorded, so

that it is possible that, if only a few stubs were present, they may have

been regarded as slightly atypical clean shanks. In one other case, boot-

ing was nearly absent. Of the 13 adults, the three males and four barred

females were alike in that the amount of booting was decidedly scanty,

being reduced to about two or three imperfect rows of rather short

feathers. The six black females were more variable, due apparently to

greater variation in length of feather rather than to variation in the

number of rows, the result being a greater variation in amount of boot.

A much larger range in the amount of booting appeared in the next

generation. The following relative grades of boot in the chicks were

- recognized: A, B, C, D, E and absent. No emphasis is to be laid on

these degrees, except in so far as they show the general distribution of

boot. A and B correspond approximately to that of the parental Lang-

shan, and C and D to that of the F, hybrids. Among the adults, not

only were there some birds heavily booted like the Langshans, some like

the hybrid and others clean-shanked like the Rocks, but one bird had two

rows of rather long feathers and one bird four rows of short feathers,

indicating that there is more than one component to boot.

126 ANNALS NEW YORK ACADEMY OF SCIENCES

TABLE I

Distribution of Booting in F, and F,.;

MUN aka B c D E | Absent} Total Remarks

3 1 i te elope ome 0 6 | 29 | F, females from 1 x F,

male from 2.

da 0 10 2 8 3 4 27 | F, females from 2 xX litter

brother.

4b 0 6 3 4 5 3 21 | F, females from 2 X re-

| ciprocal litter brother.

0 0 0 0 2 6 8 | F, females from 2 x Rock

male.

6 0 a 10 il Z) 0 30 | F, females from 1 x Lang-

shan male.

7 0 0 0 3 4 3 10 | Female Rocks X male

| from 1.

8 3 13 4 10 0 20) 30 | Female Langshans X

male from 1.

TaBLeE II

Expectation

; Observed

Ce rg rege oa Total Remarks

clean eo Sy

Clean | Booted| Clean | Booted

i F, 0 0 all 2 24 26 | Langshan females x Rock

male.

2 4 0 0 all 0 32 32 | Rock femalesx Langshan

male.

3 >» | 18.75] 5.4 | 23.6 6 23 29 | Females from 1 X male

from 2.

4a & b) F, 125 6. 42. a 41 48 | Females from 2 Xx males

from both 1 and 2.

5 F,., | 00 4, 4 6 2 8 | Females from 2 X Rock

male.

6 ee Pe O 0 all 0 30 30 | Females from 1 X Lang-

shan male.

7 ies Olea onto amoncoll lic i 10 | Female Rocks, male from

Suni wenn | CO 0 all 0 30 30 | Female Langshans, male

from 1.

The results are in entire agreement with Davenport’s and confirm his

theory of an inhibitor. The back matings suggest that the amount of

boot varies with the increase or decrease in the amount of booted “blood”.

There are, however, one or two other theoretical ways of accounting for

the observed facts. If we assume that booting is common to both Lang-

shans and Rocks and is recessive to a pair of complementary factors, both

4See above in text.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 127

of which must be present and one of which must be duplex in order to

bring about a complete suppression of the booting, the outcome approxi-

mates the observed ratios of booted to non-booted.

By assuming that the factor which exerts its effect in either the duplex

or simplex condition is sex-linked, the results shown in Table II are

obtained. The distribution of the sexes is not given, because the numbers

available are inadequate for the solution of a problem as complex as the

present one. While the correspondence between theory and observation

in this case is close, an attempt to apply it to Davenport’s data resulted

in only partial success. This may mean only that more or different fac-

tors are involved in the production of boot in Brahmas, Cochins and

Silkies than in Langshans, or that the factors causing the inhibition of

boot development in Plymouth Rocks are different from those of Tosa,

Minorca and other smooth-shanked birds used by Davenport. Among

possible factors concerned in boot production should be included those

general factors which affect feather growth, in the same way as barring

or other color factors control the color of the feathers of the boot as well

as those of the body.

Down Colors.—The Langshan chick is black dorsally but yellowish

white beneath and has white wing tips. The white ventral area often

extends upwards, particularly on the head, so that in some cases in this

Tegion only the crown and nape remain black. The white area of the

wing tips at the same time increases in size, so that the black dorsal

surface becomes reduced in amount.

The Rock chick, however, though black dorsally except for the gray

occipital spot, is usually dark gray beneath, but very often there are

several light gray or white areas, which occasionally become more or less

confluent, and in extreme cases most of the ventral surface is white and

to a limited degree overlaps the Langshan type.

In classifying the chicks, all were called “black,” 7. e., of Langshan

type, in which at most the breast region was partly pigmented. This

region in the Barred Rock chick is the last to lose pigment. All others

were classified as “barred”. While this mode of treatment proved to be

inadequate for the entire solution of the inter-relations of these charac-

ters, 1t was found, first, that both types appear in F,, but that the

“blacks” are far more numerous than the “barreds” ; second, that “blacks” |

F, interbred or backmated throw some “barreds”, but not in simple Men-

delian proportions.

128 ANNALS NEW YORK ACADEMY OF SCIENCES

CROSSES BETWEEN AMERICAN DOMINIQUE FEMALES AND LANGSHAN

MALES

Parent Generation.—Both the hens and the cock were purchased from

breeders of these strains.> The one peculiarity calling for notice is the

occasional occurrence in the Dominique hens of black or partly black

feathers (Plate XIX, figs. 6, d, e). One of the four hens used had sev-

eral such feathers. The other hens were free from them. The American

Dominiques have barred feathers (Plate XIX, figs. a, c), essentially like

those of Plymouth Rocks.

F, Generation from Langshan 6 by Dominique 2 —About 15 offspring

were reared; the hens were black and the cockerels barred. Of these, five

hens and two cocks were bred from. The black hens were like the father

as to color; the males were barred like the mother, except that a large

number of black feathers were present—some feathers entirely black

(Plate XIX, figs. r and ¢) and others barred and black (Plate XIX,

figs. p, q, S, WU).

F', Generation.—In the second generation, there were recorded 15

blacks and 14 barred birds. Three of the latter died or were killed by

animals. Of the remaining, there were 11 male and 15 females tabulated

as to color as follows: |

2 }

Barred eo e250 Baa eee ea eee 8 4

Blackie ko ee eet eae 7 7

The barred birds were fairly uniform. They were kept for about two

months, when their feathers were well developed. A few birds were dis-

tinctly darker than the rest, and one bird was much lighter. Certain

details regarding white feathers in the wings will be spoken of later.

Back Cross of F, & (Barred) to Dominique 2 .—One of the sons was

crossed to the four hens that had produced his generation. A first census

of the offspring, when the birds were small, gave 19 barred and 4 black —

birds. A later count when the birds were older gave 14 barred and 4

black. Five barred birds had disappeared. The distribution of color

and sex of 16 of these birds was as follows:

IBETVed ss eas ohne ee ee eee 7 5

BLA GI eit e eo creiaed oon ee ee nee 4 0)

5 The Dominiques came from W. H. Davenport, Colrain, Mass. ‘The source of the

Langshans is given on page 115.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 129

There were also two barred birds whose sex was omitted by mistake

in the records. The expectation is three barred to one black, which is

closely realized.

Back Cross of F, Black @ to Langshan 6 .—Five black hens were

bred to a Langshan cock of the same strain but not the actual father of

these hens. Another black hen that came from a similar cross with a

Plymouth Rock was also present in the same pen, so that some of the

offspring may have come from this hen also. There were 18 black young,

of which 11 were males and 7 females. In addition, however, there was

one barred chick. Now, the black hens had been with a barred cock to

give the F, generation. They had been for three weeks with the Lang-

shan male before the eggs fertilized by the black cock were kept for

incubation. There can be little doubt that one of the spermatozoa of the

original male had carried over and produced this bird. If this case is

thrown out, the results are consistent.

Other Features of the Crosses ——The Langshans have feathered tarsus

(booted) ; the Dominiques have clean shanks. All of the F,s recorded

were booted, though not strongly. In the F, generation, there were 14

booted and 11 clean shank, distributed as follows:

Booted Clean

2 } & $

1 SIZ EIN EY6 le geeeelene eee arora 7 , 1 2

BS EC Kaa eer euateaeroctonee 1 4 5 3

It is clear that booted shanks dominate® but imperfectly in this cross,

as in other crosses of poultry. Some of the F, offspring had heavily

booted legs; others were like the F, generation. No sharp line between

the classes in the F, generation could be drawn.

There is no evidence of any association here between black and booted

(the paternal combination) and barred and clean-shanked (the maternal

combination).

When the barred and booted F, male was bred to four Dominique

hens, the results are shown in the next table:

Booted Clean

Q 3 Q i)

1 BEWTEIENG aS otaeptovotors Bree meieneLS 6 1 4

TRNSVSE callietees ane il 0 3 0

®TIn the sense that an inhibitor is present in clean shanks.

130 ANNALS NEW YORK ACADEMY OF SCIENCES

In this case, the male was heterozygous for condition of tarsus; the

hens pure and recessive. The result calls for equal distribution of booted

and clean shanks unless “association” occurs. The numbers are too small

to have any significance. Even as they stand, however, they have no

meaning, if coupling be made responsible for the distribution of the

characters.

When the Langshan male was crossed to the black hens (both sexes

booted, but the hens heterozygous) all of the offspring were booted,

which is in accordance with dominance of booted shanks. —

White Feathers in Wings.—In the F, young birds, the presence of

white and partly white feathers in the wings was noticed (Plate XIX,

figs. f-n; 0, v and w). They were most obvious in the black birds,

perhaps because of the sharper contrast. These feathers are some of the

primaries and a few of the coverts at the base of the primaries. As

shown in Plate XIX, figs. f to k, they are rarely pure white, but often

mottled or splotched. They were not recorded in the F, birds, and if

present they were overlooked. The records of birds without and with

these white feathers were as follows:

Q }$

Barred: no: whites sie see een 6 4

REA Meminer ek eo Cho orn co Gra'G d-olcu om a OG 2 6

Black, no ‘white:2232 23s. eee eee 6 Rete 4

eee (AU Ua Ter BAA Wabis,Gccud o.0 6800 a Le 3

In all, there were 20 chicks without and 6 with white feathers. This

looks like a case of Mendelian inheritance, but it may be purely a coinci-

dence.. We do not know how often such feathers occur in chicks of the

original breeds, or whether they are only juvenile, or physiological effects

of the condition of the bird. Probably they would have disappeared in

later molts, had.the chicks been kept longer.

When the Langshan cock was bred to the black F, hens, 4 of the chicks

had no white and 14 had white in the wings. If the black male is hetero-

zygous for this condition, the result is not in accordance w ith the assump-

tion that this is a Mendelian recessive character.

When the Dominique hens were bred to the F, barred males, hare

was no white in the 15 recorded offspring. This result is not in harmony

with the same supposition, but the black male used in the last experiment

was not the same father as for the barred males of the first cross. The

father of the barred male in the first case was a brother of this one. It

is still possible, therefore, that one male was homozygous and the other

heterozygous for the white-feathered condition. Without, howeyer, fuller

information, not much weight can be given to these results.

MORGAN AND GOODALE, INHERITANCE IN POULTRY 131

Color of Legs—It has been stated by Bateson and by Pearl that yel-

low and black shanks in certain breeds of poultry show “sex-linked”

inheritance. This is not apparent in the Langshan-Dominique crosses, ex-

cept in so far as black shanks accompany black color of feathers. For ex-

ample, in the F, generation, there are recorded 13 black birds with black

legs. Of these, 5 were deep yellow on the under side of the feet. In

addition, there was one male that had yellow shanks and yellow under

the feet. There were recorded 12 barred F, chicks with yellow shanks.

Of these 12 birds, 4 are recorded as having very pale yellow or whitish

legs. It would appear from this case that black and yellow shanks

accompany black and barred plumage, at least as a rule.

Tn the back cross of the F, barred male to the parent Dominique hens,

in which there were barred males and females and only black females,

all 4 of the black birds had black legs, while all 12 barred birds had

yellow or pale legs. Among these 12 barred birds, there were 5 with

pale legs; in 2 of these and in one yellow, there were spots of black or

dark color, at least on the tarsus.

These. rather meager figures, as far as they go, show that shank char-

acter and color of plumage go together, and that black shanks and yellow

shanks are only an accompaniment to sex-linked inheritance of plumage.

The data are manifestly few, however, and it may well happen that the

two characters may appear disassociated.

Color of Bill—The color of the bill seems to run a parallel course.

Full records for the back cross given above were kept. Here, 13 barred

birds had yellow bills and 5 black hens had black bills, but one of the

latter had much yellow on it, and two of the former had black: one was

black with yellow tip and the other was yellow and black. There is much

variability in the color of the bill, and the above statements are insuffi-

cient to warrant any generalizations.

THEORETICAL CONSIDERATIONS

The current formula for sex inheritance in fowls represents the female

as heterozygous for sex, F-O, and the male homozygous, O-O. If F is

identified with a special chromosome connected with sex determination,

the formula calls for one more chromosome in the female than in the

male. At present, evidence on this point is conflicting and insufficient.

It is true that Guyer has described two kinds of spermatozoa in the male,

one with an X and one without. If this X is the same as in other ani-

mals, then the spermatozoa containing it must be female producing, and

the female should contain one more chromosome than the male. This

means that the male and not the female is heterozygous for sex. The

132 ANNALS NEW YORK ACADEMY OF SCIENCES

experimental evidence is flatly opposed to this latter interpretation, and,

therefore, until Guyer’s evidence is confirmed or refuted, the case must

be left open.

On the other hand, if, as the experimental evidence shows, barring is

“repulsed” by femaleness and if both of these factors are carried by

chromosomes, the formulas are deficient in having no chromosome to

carry barring,—a contradiction of terms. It may be, however, that the

X-chromosome in fowls has a mate which we may call Y which would

carry barring but not femaleness. The formule would then be:

NOTA Oxerc tere crsccneheee eae scr ne oer A eee xX —Y

JY RET) SRA nS ee A aren hee gar by ee Y— Y

XY Female

YY Male.

On this interpretation, the factor for femaleness would be contained

in X but absent from Y, while barring is contained in Y. ‘This scheme

is compatible with the experimental evidence and gives consistent results

for all combinations.

The irregularities that have been observed in the “reduction division”

both in birds and in man suggest the possibility that the sex chromo-

somes are united to other chromosomes as in some other animals. If the

union is variable, as in the nematodes, it may be that the X and the Y

(if Y exists) may sometimes pass to the poles of the spindle during

reduction in conjunction with other chromosomes and sometimes be free

to pass to the poles independently. If further study should establish

this view, it will have a very direct bearing on the relations discussed

above. If the factor F for femaleness is carried by chromosomes attached

to one member of another pair, the mate of this member may be the

chromosome that carries the factor for barring. If this were the case,

however, interchange between these two members would lead to the

barring factor being transferred to the chromosome attached to the sex

chromosome. This is in contradiction to the experimental evidence

which would lead rather to the conclusion that a Y element lacking

the factor for barring is present. The Y may be attached to the mate

of the chromosome carrying the sex factor.

At present, only a few cases have been discovered in which a sex-linked

character is dominant, viz.: in fowls and in one character in Drosophila.

The only other cases, besides the one in poultry in which sex-linked in-

heritance occurs and sex is heterozygous in the females, is that of Abraxas

and that of canaries. In both of the latter, the sex-linked factor is re-

MORGAN AND GOODALE, INHERITANCE IN POULTRY 133

cessive. There are no a priori grounds why a character of this sort may

not be dominant, if some other Mendelian characters may also be domi-

nant.

The factor for black, N, is treated in our formule as present in all of

the gametes both of the female and of the male. It is not allelomorphic

to barring, B, although its presence in the female-producing egg when

barring is present in the correlated male-producing egg may appear to

bear this interpretation. From the chromosome point of view black may

be, so far as we know, in other chromosomes than those carrying barring ;

hence its more general distribution.

BIBLIOGRAPHY

American Poultry Association, The American Standard of Perfection. 1905.

Bateson, W.: Facts Limiting the Theory of Heredity. Science, N. S., Vol.

XXVI. 1907.

: Mendel’s Principles of Heredity. 1909.

: The Inheritance of the Peculiar Pigmentation of the Silky Fowl.

Jour. Genet., Vol. I. 1911.

Bateson, W., and PuNNeETT, R. C.: The Heredity of Sex. Science, N. S., Vol.

XXVII. 1908.

Brown, H.: Races of Domestic Poultry. London. 1906.

DAVENPORT, C. B.: Inheritance in Poultry. Carnegie Pub. No. 52. 1906.

: Inheritance of Characteristics in Domestic Fowl. Carnegie Pub. No.

121. 1909.

GoopALE, H. D.: Sex and its Relation to the Barring Factor in Poultry.

Science, N. S., Vol. XXIX. 1909.

: Breeding Hxperiments in Poultry. Proc. Soc. Exp. Biol. and Med.

Vol. VII. 1910.

Hanviey, P. B.: Sex-limited Inheritance. Science, N. S., Vol. XXXII. 1910.

HaGrepoorn, A. L.: Mendelian Inheritance of Sex. Archiv. Ent. Org., Bd.

XXVIII. 1909.

PEARL, R., and Surrace, F. M.: Studies on Hybrid Poultry. An. Rep. Maine

Agric. Exp. Station. 1910.

: On the Inheritance of the Barred Color Pattern in Poultry. Archiv.

Ent. Org., Bd. XXX. 1910.

: Further Data Regarding the Sex-limited Inheritance of the Barred

Color Pattern in Poultry. Science, N. S., Vol. XXXII. 1910.

SPILLMAN, W. J.: Spurious Allelomorphism; Results of Some Recent Investi-

gations. Am. Nat., Vol. XLII. 1908.

STuRTEVANT, A. H.: Another Sex-limited Character in Fowls. Science, N. S.,

Vol. XXXIII. 1911.

TEGETMEIER, W. B.: The Poultry Book, etc. London. 1867.

WaicutT, L.: The New Book of Poultry. London. 1902.

PLATE XVII

ED PLYMOUTH ROCKS, LANGSHANS AND THEIR CROSS-BRED OFFSPRIW

iis a Tae NaC

‘1. Barred Plymouth ‘Rock cock. —

2. One of the parental Langshan hens.

8. Langshan cock. ‘Stock of Mr. Ives.

4, One of the parental barred hens. This particular hen is

ey White Rock male X Barred Rock female.

_ Fes. 5 and 6. The F, from Barred Rock cock by Langshan hen.

A Figs. 7 and 8. The F, from Langshan cock by Barred Rock hen.

> sedi) ’ = : i)

RGN 1O (iie-aaod9

eel oa aig He sod sLblaothed ate Ae

joleina? vba halves x ‘hkera ‘ase

sit coy al A doo ood ode ago mn al

ANNALS N. Y. Acap. Sct.

VOLUME XXII, Pharm XVII

PLATE XVIII

FEATHERS OF LANGSHANS, BARRED PLYMOUTH sacs AND THEIR Se

‘ " Hackle feathers (except 11 and 14) ‘from the various types. of pirde used i in

Best ny _the experiments.

Basa and 4. fou a pure bred Barred Rock female.

‘Fie. 5. From a second Barred Rock female. Note the differences in the Bie

ness of the barring. Cg

6. From the hen shown in Plate XVII, Fig. 4. i et

7. From F, barred female. — (

Fig. 8. From a pure bred Barred Rock cock. . j

9. From same bird, illustrating a partially black feather occurring in

pure bred stock.

Fies. 10, 11, 12, 14. From F, bird shown in Plate XVII, Fig. ay,

Fig. 10. Hackle feather.

Fie. 11. Breast feather.

Fig. 12. Shows the Jungle fowl coloration.

Fie. 13. From bird shown in Plate XVII, Fig. 7.

Wie. 14. Primary, to show reduction in barring.

Lig fe Rea ees ®

PUIG Ah. GEA Get ra EURUNAME it canta an

tii Oem wbittd ths ang) evoliny, off mort CM Dane i sao)

minus Orel a Beate

Jtaenet Aooh base bene hla th ‘aie

ve GAs Te eopusto Dib sii shod aisinel obo05 alae bine 4s

ie iL ‘warritioo0 fed deo} oat wise i gute baie Summa s

Bai AYA aks 14 ua vie bald 1 port bs ’

bis rain :

siniteiotoy | ‘wor yen gilt

Pie ae ohn ths Ge ies gah bata

”

PLATE XIX

FEATHERS OF AMERICAN DOMINIQUE FOWL

OLB ,

a. Dominique hen. Barred feather.

b. Dominique hen. Black feather. /

. €. Dominique hen. Barred feather.

ee Domini me hen. Black feather.

Black feather.

ae " feathers.

Figs. in, Three white covert feathers from mines

me ‘ oped feathers at their base.

Fic. w. ee of barred F,, showing a nearly

barring: on some other feathers.

Zeb fe VLE

OFT VLPeoey, OR: AR OUS >» oh

Sat peng a

oii ior BOe: ithe ods cat ballet ‘he aarunlt sq

ae gatwods etiur parent ie ass arioge

Neehteb ae’ uel Bara 29 mea ‘oy sie aed Bore

Sppligead il Pinter HGH CE BPI iE hinon o

VOLUME AATI, PLATH AIX

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THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Naruran History, 1817-1876)

OFFIcERS, 1912

President—EMERSON McMixuin, 40 Wall Street

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[ANNALS N. Y. Acap. Sct., Vol. XXII, pp. 155-160, Pll. XX-XXII, 25 July, 1912]

ON THE DICTYONEMA-FAUNA OF NAVY ISLAND, NEW

BRUNSWICK

By F. F. HAHN

- (Presented by title before the Academy, 13 May, 1912)

CONTENTS

Page

IGDEFO CLC THIGIN ey nee ered alten Gididle a-cra Oc. & choiena: cor ORI crete RCC a neers maces eer uere 135

Gury. WBA UNO YS BI PB igs 6 Sinton pis a oo charg oo 6 Heels Se eC Ick hs cat rcs 136

Astogeny of Dictyonema flabelliforme in comparison with that of Stawro-

GirUPUUSs. 05.6504 60 6000 OOOO UID b O00.0'0 0.0 00-00 0100 6 CO CICIEIG DOING Olte aiCeutClc aD Ero 144

On the structure of Dictyonema flabelliforMme... 2.1.2 cece cece cee eee cee 146

Mode of life of Dictyonema flabellifOrMme. ... 0... cece cece eee ee eee eens 148

On the occurrence of stratigraphical range of the varieties of Dictyonema

MOGI ORTIGsias ce Sooceidoatadcuesucdcobe 0 oOo COC Ud uc oUOO oO coo COCO CUO 151

Significance of the varietal range and its phylogenetic value............. 153.

INTRODUCTION

One of the older modes of paleontological investigation, the study of

the range and significance of variability, seems now to be somewhat

overtaken by and neglected on account of the excellent results obtained

in the study of the stages of earliest youth, growth and senescence. My

own studies on Jurassic invertebrates, which made me familiar with a

vast number of individuals of many species, brought me to the convic-

tion that a combination of these two modes of treatment would, in every

case where the given material is sufficient, lead to an advance greater

than would be possible by either method alone. Thus, when I worked

over in the Museum of Columbia University the wonderful collections

from the Dictyonema-shales made by G. van Ingen and W. D. Matthew

at Navy Island, N. B., from Division III, b, ec, commonly regarded as

Upper Cambrian,* of course my first thought was to see whether I could

find any relationship between the range of variability and the history of

the races among the Dictyonemas. And this animal association seemed

well adapted to this kind of study, for though the Dictyonema-shales

contain a vast number of individuals, there are only four species com-

mon, two of which are graptolites.

1 According to the trilobites found in Sweden and BHngland, it must apparently be

placed at the base of Ordovician.

(135)

138 ANNALS NEW YORK ACADEMY OF SCIENCES

Description: Generally infundibuliform, branches more or less parallel, fre-

quently dividing, width of tissue 9-10 mm., cross-threads somewhat irregular,

of medium number; open rectangular meshes; thecze 15 to 17, projecting, of

medium size.

Notes: Slight changes of the characters are fully shown on the diagram,

Plate XX. Greatest length observed about 200 mm., width 190 mm.; approxi-

mately 2x 150 branches.

The variation of the number of thece is marked, 16 being present on

the American examples, but only 10-15 in 10 mm. on the European ex-

amples. This fails to be of general value, as G. F. Matthew has pointed

out, since the figures of Brégger (5) and Tullberg (8) also exhibited 16

thece and Westergard states recently a usual number of 15-16 thecz on

the Swedish examples ((4), p. 58). When comparing the form, col-

lected by Prof. A. W. Grabau from Skane (region of Fogelsang), with

the American specimens, a more delicate character of the whole tissue of

the European type is apparently the one observable difference on which

a separation of var. acadica Matth. and “forma typica’ of the Swedish

authors can be based.

Var. conferta Linrs (ms.)™

Apotypes :

(6) 1860. GOpprrRT, pl. 36, fig. 11, 4.

(9) 1861. Satrer, Geol. Survey Gr. Brit., Mem. 3, pl. 4, fig. 1.

1881. SALTER, Geol. Survey Gr. Brit., sec. ed., p. 535, pl. 41, figs. 1, la, 1b.

($) 1882. TuLteerre, p. 20, pl. 3, fig. 3 (1, 22).

(4) 1909. WESTERGARD, pl. 3, fig. 7.

Description: Characterized by Brégger ( (5), p.36) as having very fine and

close network and cross-threads of the same kind.

Matthew” added “commonly vasiform, cross-threads more frequent than in

acadica (5-7). 15-17 thece are usually met with in 10 mm.”

I have observed even 18 on a typical specimen ; young specimens often

show only 14-15 thece. The rigid aspect of the branches and their

regular branching, which commonly takes place in the same level, seem

rather important additional features. Here, too, a little more delicate

structure of the whole tissue distinguishes the Swedish specimens, so that

Westergard found 13 branches in 10 mm. of width, while 8-10 branches

may be usually numbered on the American variety. The largest ob-

served colony measures 22 mm. in width, occupied by 19 branches, and

82 mm. in length.

ei Broégger (5) furnished as long ago as 1882 an undoubted description of this form,

while Pocta created his species “conferta’ not earlier than 1894 (Systéme silurien,

Bryozoaires, ete., Vol. WIII, t. ler, p. 194). The last ought to be renamed, if this is

necessary at all.

2 Loe. cit.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, VN. B. 139,

Var. norwegica Kjerullf.

Apotypes:

(7) 1865. Ksrrur, p. 2, fig. 4 (1-3).

(2) 1906. Mosere, pl. 1, fig. 7.

(4) 1909. WesrercArp, pl. 30, figs. 8, 9.

Description: After Broégger ( (5), p.36), thick cross-threads, close network ;

small, short, rounded meshes.

I observed only 15 thece. Young stages like var. acadica. Crossings

of all kinds into true acadica occur. A shght distinction between my

specimens and the Swedish ones is in so far recognizable as the first are

furnished with a little more elongated meshes, but they can be precisely

compared with Westergird’s figures. My largest colony is 40 by 40 mm.,

while Westergard cited a specimen 115 by 30 mm. in size.

Var. ruedemanni nom. nov.

Typical figures:

(1) 1904. R. RuzpEMANN, pl. 1, figs. 16-20, 22 (12-15, non fig. 21).

Apotypes:

(10) 1895. G. F. MatrHew, Trans. N. Y. Acad. Sci., Vol. 14, pl. 49, figs. 1, 2.

(The figures are not quite correct, showing a too open tissue. )

Compare

(5) 1882. BroeeEr, pl. 12, fig. 19.

Description: In nepiastiec stage strongly branching at an almost acute angle.

Full-grown specimens with crowded, more or less parallel branehes up to 14 in

10 mm. width. Dissepiments appearing early, but afterwards irregularly scat-

tered. Thece small, not much projecting. Adhesion organs relatively common.

Notes: Generally funnelform, sometimes of intermediate shape (pyriform)

towards the vasiform variety “conferta.” Measurement of a typical pyriform

specimen :

Length Width in mm.

10 10

20 25

25 35

48 65

The specimens obtained from Navy Island (N. B.) fully agree with

Ruedemann’s figures, but several specimens considerably surpass the

given characters in the same direction. In Europe it is rare, but abun-

dant in America, at Navy Island as well as at Schaghticoke. Largest

measured colony 165 mm. in length and 140 mm. in width.

Var. desmograptoidea var. nov.

Description: Infundibuliform. Open meshwork of irregular, mostly sub-

oval fenestrules. Branches undulating and sometimes coalescent. Dissepi-

140 ANNALS NEW YORK ACADEMY OF SCIENCES

ments quite irregular, partly thickened. Thece 15 in 10 mm., not much pro-

jecting, rarely visible. ;

Note: Sometimes an old stage of var. acadica has these characters.

J. Hall’s!® Dictyonema irregularis resembles most this variety, but

has a closer arrangement of the branches (25-28 in 1 inch) and occurs

in higher Ordovician of Canada and Great Britain. Dictyonema hom-

frayi, described by T. Hopkinson and Ch. Lapworth,* seems to be a

closely related species. Desmograptus intricatus, as shown in fig. 30, p.

609, by Ruedemann (1), exhibits exactly the same type in early stages,

though belonging to the Chazy and having appressed thece. No well-

preserved, large specimens are known, but several pieces are found indi-

cating a width of the colony of more than 60 mm.

Staurograptus dichotomus Emmons var. apertus Rued.

(1) 1904. R. RUEDEMANN, pp. 612-614, pl. 2.

(11) 1891. G. F. Marruew, “Bryograptus kjerulfi,’ Trans. Roy. Soe. Canada,

Sect. IV, No. VI, p. 35.

1892. G. F. Matruew, “Bryograptus patens Matth.,”’ No. VII, p. 95, figs.

la, 1d (excl. lower figure of 16).

(10) 1895. G. F. MatrHew, id., p. 268, pl. 48, fig. 4 (?).

1895. G. F. MatrrHew, “Bryograptus lentus Matth.,” id., p. 270, pl. 48,

fig. 2.

1895. G. EF. Matruew, “Clonograptus proximatus Matth.,” id., p. 265, pl.

48, fig. 1.

In 1895, G. F. Matthew tried to keep separate five species of Clono-

graptus and Bryograptus, occurring in Division 3, band ¢c on Navy Island,

N. B. In 1903, Ruedemann’ followed him, believing that four of

Matthew’s species were again recognizable, but a year later, in his Mono-

graph on the Graptolites, he came to the conclusion that practically all

these species and even genera present only definite stages of growth and

preservation of Staurograptus dichotomus, a species established by Em-

mons in 1855, but rather dubious up to 1904. Ruedemann, indeed,

furnished such a description and figures of this species, especially of the

variety “aperta,’ that complete identity with the species of Matthew

enumerated above becomes apparent, if we look over the types of

Matthew, which fortunately were in part accessible to me.

18 Canadian Organic Remains. Geol. Sury. Can., Dec. 2, p. 136, pl. 20, figs. 1, 2.

1868.

144 Graptolites of the Arenig and Llandeilo rocks of St. David's. Quart. Journ. Geol.

Soc. London, Vol. 31, p. 668, pl. 36, fig. 13. 1875.

“The Cambric Dictyonema fauna in the Slate Belt of Eastern New York.” N. Y.

St. Pal. Ann. Rep., Bull. 69, p. 938.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 141

The type specimen of Clonograptus proaimatus (1895, pl. 48, Figs.

la, 1d) proves to be almost identical with Ruedemann’s figure 23 on

plate 2, besides a slightly greater elongation of the thece. Of these, I

have found distinctly 8 in 8 mm. of length of the branches, the latter

not surpassing 25 in number. The fork-lke mode of branching seems

highly characteristic.

On the same slab, the type of Bryograptus lentus Matth. (1895, pl.

48, fig. 2) is found, appearing at first glance very distinct from that of

“Clonograptus proximatus ;” but here, too, 9 to 10 thece of a breadth up

to .75 mm. are seen, showing the same aspect as Ruedemann’s figs. 19

and 21. Moreover, this slab exhibits another crowd of branches which

in part are completely identical with “Clonograptus proximatus,” while

a few branches turned to side view show the real aspect of “Bryograptus

lentus.” Here again, 9 thece are recognizable in 10 mm. of length of a

branch which is 1.6 mm. thick and of the same shape as Ruedemann’s fig.

15. This group of Staurograptus has still greater importance, since three

individuals of medium sizé have their sicule preserved and present the

characteristic side view of Ruedemann’s fig. 6. As a great many other

specimens of the collection which formerly were determined as “Bryo-

' graptus patens,’? and indeed readily conform to Matthew’s fig. 4, on

plate 48, really belong to nepiastic stages of Dictyonema,'® a similar

reference of the type of Matthew seems quite possible, but I have not

seen the type specimen, and thus this explanation may still be ques-

tionable.

A few additional notes can be added to the description by Matthew

and Ruedemann.

The preservation showing the thece in full size and the sicula is rare

on examples surpassing a diameter of 5 mm. The thece, scarcely more

or less than 9 in 10 mm. length of the branch, are sometimes irregularly

exposed, 6 smaller ones being visible in 4 mm. of the distal end and 4 a

little coarser ones in 4 mm. of the proximal part of the same branch.

Young stages, especially when crowded together, are almost inseparable

from those of Dictyonema flabelliforme var. ruedemanni, while the bryo-

graptoid neanastic stages of D. flabelliforme var. acadica and conferta

are easily mistaken for the “Bryograptus lentus” aspect of laterally com-

pressed Staurograptus. The great thinness of the periderm of Stawro-

graptus, which never shows such a tubulose structure or wrinkling as in

the case of Dictyonema (compare the above), is a very remarkable fact

and helpful in distinguishing these two genera. In the same respect, the

16 Compare Westergard (4), pl. 3, figs. 5 and 6.

142 ANNALS NEW YORK ACADEMY OF SCIENCES

more rigid characters of the mucronate apertural margin of the thece in

Staurograptus may be noticeable.

Unfortunately, I was not successful in finding any remains which can

be compared with “Bryograptus spinosus Matth.” ((10), p. 269, pl. 48,

fig. 3) and “Clonograptus spinosus Matth.” ((11), p. 9%, pl. 7, fig. 2),

which is said to present only 8 thece in 10 mm. of length of the branch

and a distinct “axis or virgula.” Nevertheless, here, too, the identity

with Stauwrograptus dichotomus seems to me not quite impossible. Nor

-could I observe any specimens belonging to “Bryograptus ? retroflexus”

(Matthew (10), p. 271) or to Callograptus (1. ¢c., pl. 48, fig. 5), which

therefore must be at least extremely rare.

On the contrary, with an abundance of individuals there is found

Monobolina refulgens Matthew

(11) 1891. G. F. MatrHew, p. 44, pl. 12, fig. 6.

(12) 19038. G. F. MattrHew, Report on the Cambrian Rocks of Cape Breton.

Geol. Surv. of Canada, Ottawa, p. 210, pl. 11, fig. 4, pl. 16, fig. 2.

Since generally separated valves and even broken pieces occur, nothing

can be added to the careful description of the author, besides the obser-

vation that this species is seen on the same surface associated with all

varieties of Dictyonema flabelliforme and with Stawrograptus dichoto-

MUS.

Lingulella nicholsoni (?)

Almost the same is true of a linguloid shell, cited by Matthew in

1895 (p. 273) as Lingulella nicholsoni (?) and in 1903 ((12), p. 204)

doubtfully called Lingulella “lepis Salter.” The specimens of the Co-

lumbia collection usually have a length of 4 to 5.5 mm., a width of 3 to

4.8 mm. and exhibit a distinct ridge on the interior of the pedicle valve.

They differ from the English type as described by Salter ((9), second

ed., p. 538, fig. 11) in a relatively greater feebleness and scarcity of the

lines of growth. It may be of interest to note that a very similar

“Lingulella” of Scandinavia is referred by Brogger ((5), p. 44, pl. 10,

fig. 5, from the Tremadoc shales) and by Moberg ((2), from the Bryo-

graptus zone) to L. lepis Salter.

In addition to this detailed discussion of the Dictyonema fauna, a

callograptoid graptolite of the upper Beekmantownian may be described,

because of its interesting phyletic relationships.

Callograptus grabaui sp. nov.

Flabelliform or shrub-like, not more than 15 mm. in length and 10 mm. in

width of the dendromes observed. Short, non-celluliferous, basal stem (1 mm.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 143

long, .5 mm. wide) with terminal expansion to an adhesive bulb (of .8 mm. in

width). Branches 10 to 13 in 10 mm. width, .3-.4 mm. wide, closely arranged,

sub-parallel, sometimes slightly flexuose. Sicula of one specimen 7 mm. long,

determinable as being a very thin tube about 1 mm. in length, with a minute

basal disk .5 mm. in width. Within the first 5mm.

of the length of the dendrome, a very frequent

branching, sometimes of monopodial aspect, with

a common angle of 50-60° takes place. Branches

of the third and fourth order are usual, those of

the sixth still observable. The interspaces be-

tween the branches vary from .83mm. to 1 mm. in

width. Most of the specimens are dorsally de- BiG, i—<Collegeniins acta

pressed and therefore, as commonly found in sandy sp. nov. (3/1)

shales, they do not exhibit the thece. Only a few

branches, laterally compressed, show 19 to 21 thece in 10 mm. of length of the

branch, as sharp spinelike prolongations, quite similar to those of Dictyonema

flabelliforme (cf. Ruedemann (1), p. 601, fig. 26). Very seldom one or the

other of these apertural processes reaches the neighboring branch, joining it

like a dissepiment. The periderm seems to be composed of two layers, of

which the greenish external one adheres to one side of a split slab, while the

black interior layer remains on the other side, distinguished by a wrinkled sur-

face. Aside from this no striation is visible.

Stratigraphic Position and Mode of Occurrence: The type specimen

was sent to the Columbia Collection by Victor Ziegler, who obtained the

fossil from Center County, Pa., “2000 feet from the bottom of the lime-

stone series which are referred by Collie to the Beekmantown.” 7 The

animal mass is profusely scattered on the sandy surface of the thin-

bedded, impure limestone.

The close relationship of this species to Callograptus salteri Hall and

C. compactus Walcott is proved by the following table; it is contrasted

only by the distinction of the thece.

C. saltert C. compactus C. grabaui

Max. observed size........ 17 mm. 40 mm. 15 mm.

Number of branches in

10 mm. width ........ \ Le te ‘ ts

Thickness of branches..... .) mm. .o--4 mm. .o- 4 mm.

Number of theez in 10 mm. 14-18 12-14 19-21

Thickness | 2 X thickness of | Average thickness

Interspaces ............. | of branches. branches. of branches.

17This zone may belong to Ulrich’s Stonehenge limestone.

144 ANNALS NEW YORK ACADEMY OF SCIENCES

J. Hall, who first described Callograptus, laid special emphasis on the

characters of the whole dendrome and as in this respect the new species,

C. grabawi fully agrees, it must be for the present referred to this genus,

even though the deviation in the aspect of the thecee shows two divergent

lines.of evolution among this “genus,” similar to those in Dendrograptus,

Ptilograptus ete. The striation of the rhabdomes, which some authors

regard as an important feature of the genus, is not mentioned in the

early description of Callograptus salteri by J. Hall, while Ruedemann

observed such longitudinal striations “when the thecal tubes have been

pressed through the periderm.” ‘This is also entirely the case in the new

species.

There are three other genera of the Dendroidea which hold a median

position between Dictyonema and Dendrograptus and to which C. gra-

baui could be considered to belong. Odontocaulis is of quite the same

shape, but occurs a little later in Ordovician and Silurian time and

differs in its celluliferous stem. Calyptograptus, a Niagaran genus,

shows independence of the main rhabdomes down to the root as an essen-

tial feature. Rhizograptus, in its genotype (the Niagaran “bulbosus’),

has branches, more or less reticulated, joined or overlain by others. The

dissepiments of C. grabawi are extremely rare, so that a position among

the true Dictyonema would be erroneous, likewise, a union with Dendro-

graptus seems to transgress the natural and originally assumed limits of

this genus, considering the presence of cross-bars and the shrub-like, or

perhaps even funnel-form growth of the polyparium.

ASTOGENY OF DICTYONEMA FLABELLIFORME IN COMPARISON WITH THAT

OF STAUROGRAPTUS

The collection of Columbia University contains such a number of the

earlier stages of the two genera that I was easily able to select a com-

plete series beginning with the sicula and closing with the full-grown

colony and this series is now preserved in the paleontological museum.

Still, only a few remarks can be added to the illuminating description of

Ruedemann so far as the later stages of D. flabelluforme, as described by

this author, deviate in some respects from the normal ones presented by

“forma typica.” All essential features, however, fully harmonize with

Ruedemann’s observations.

After the formation of the sicula, marked by a short, stout initial part,

on Dictyonema, by a slender, curved one on Stawrograptus, the katem-

bryastic’® series begins with the budding of the first, second and third

18H. R. CUMINGS: “Development of some Paleozoic Bryozoa,” Am. Journ. Science,

Vol. XVII, p. 50. 1904.

eee

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 145.

thece. ‘These nepiastic stages of Dictyonema are characterized by the

large angle of divergence of the first theca, the next following tripod

aspect, particularly by the 4-branch-stage fairly symmetrical and basket-

like (compare figures on plate XXI), while Stawrograptus has the first

bud apically derived and mostly appressed and passes afterwards through

a characteristic stage with six branchlets resting in a plane vertical to

the axis of the sicula. It further exhibits a slight asymmetry because of

accelerated divisions of one of the primary branches. The following

neanastic stage develops not only wholly the specific features, but even

some of varietal value, while nemas become rare. In D. flabelliforme

var. acadica, a subsequent dividing of the co-equal four primary branches

happens after this, though on one or the other retarded branch 3, 4 or

even 6 thece may form a uniserial twig before its division. (Compare

figure 2 on Plate X XI.)

The first dissepiment, being the last appearing generic feature, is de-

veloped as a rule when branches of the third or fourth order are present..

Thus very often a lateral compression of the movable branches will

furnish a striking bryograptoid view. The same is true in var. conferta,

besides the fact that with the latter a monopodial dividing frequently

occurs. The neanastic stages of var. rwedemanni, however, are charac-

terized by irregularly crowded branching and early appearance of cross-

threads, as fully described and figured ((1), plate 1, figs. 13 to 19) by

Ruedemann. Beyond that, several of the neanastic specimens of this.

variety prove to be so much accelerated, that immediately from the sicula

a thick scarcely resolvable bush of rhabdomes appears to arise in a real

dendrograptoid manner of growth. Neanastic stages of var. norwegica

and var. desmograptoidea do not deviate from those of the forma typica

except in one specimen of the latter variety, which has the irregular

division of the branches at a remarkably large angle somewhat acceler-

ated.

Staurograptus dichotomus generally runs through a 5- or %-branch-

stage and the individuals of this collection turn into var. aperta by slow

or retarded dividing of an angle of 70° to 90°.

The ephebastic stage of Dictyonema flabelliforme chiefly establishes.

the characteristic outline of the colony which is infundibutiform in var.

acadica, norwegica, ruedemanni and desmograptoidea, pyriform in ruede-

manm m. tf. conferta and vasiform in conferta. It is in this stage that

the essential features of variety desmograptoidea and norwegica are fully

developed. Basal expansions are now more commonly found. The ephe-

bastic stage of Stawrograptus, relatively rare, regarding the surplus of

younger individuals, shows a diminution of the angle of divergence, en-

146 ANNALS NEW YORK ACADEMY OF SCIENCES

largement of the thece and an oblique tendency of the unsupported rhab-

domes, which leads to a bryograptoid exterior.

Gerontastic features are noticeable in some specimens of Dictyonema

flabelliforme var. conferta, in which the development of the main varietal

feature is carried to an extreme as exhibiting a reduction of the diameter

of the synhabdome on its posterior end. Very large individuals of var.

acadica indicate an undulating shape of the rhabdome, while this is tele-

scoped in other specimens into ephebastic and even neanastic stages.

As to the possibility of keeping distinct the young stages of Dictyo-

nema and Staurograptus, the 4-branch-stages of the first one, which does

not go beyond the declined’® position, and the 6-branch-stage of Stawro-

graptus, which gets into a deflexed or even horizontal position of the

branches, are of greatest importance. A similar difference holds true in

regard to Bryograptus, of which Westergard has recently described beau-—

tiful specimens of B. hunnebergensis ((4), pl. 5, figs. 10-23).

I may call attention to the fact that the Dictyonemas here found ex-

hibit all features of a true epacmic genus, since this increasing period of

evolution is indicated by (1) the relatively long duration and the clear

distinction of the early stages, because of non-existence or at least only

the beginning of acceleration of the characters; (2) the appearance of

new “postspecific” features in the metephebastic age, which may become

inherent ones and may be pushed back by acceleration in the descend-

ants; (3) the fact that senile features make their appearance late and

are rare, if we confine them to those which doubtless manifest a decline

of the individual or the colony; (4) the fact that the genus in question

maintains its position at the base of a more or less widely branching

evolutional series. All these features are shown in Dictyonema flabelli-

forme from Navy Island, N. B.

ON THE STRUCTURE OF DICTYONEMA FLABELLIFORME

Though the specimens available for study were compressed upon the

shale and so do not permit investigations similar to those made by

Wiman, I have nevertheless been compelled to direct my attention to the

points raised by these investigations, since several recent writers have

spoken rather slightingly of them or have considered them open to

question.

At first glance, a striking difference seems to exist between the grapto-

litic mass, one rhabdome exhibiting a reflecting silvery surface while

others show a matt blackish one. By cautiously scraping on the best-.

19 See Ruedemann (1), p. 485.

ati tl tod

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 147

preserved specimens, the reflecting layer is found beneath the matt ex-

terior, which usually adheres to one side of the split slab, the reflecting

layer remaining on the counter side. If we shave the reflecting mass

away, again a matt black skin can be recognized. This is apparently the

same type of observation made by Frech on the graptolites of Sadewitz.*°

He states that a dark horny cover is separable from a brilliant interior

layer and that the latter consists of calcite crystals. I could not get, it

is true, any reaction on touching the mass with hydrochloric acid.

Nevertheless, I believe that the two sheets described may represent the

epidermal and mesodermal tissues of these ancient hydroid zoaria.

As to the composite character of the rhabdomes, on all better pre-

served branches, the wrinkled structure long ago recognized by a great

many authors can easily be seen in var. acadica and conferta, while rarely

observable in the other varieties. After much search, I succeeded in

finding a few dendromes on which pyritization had partly taken place, so

that several minute tubes about .10 mm. in width are shown in relief

running along the thece in a slight curve (compare Ruedemann (1), p.

607, text figure 28) toward the aperture.

The terminal pores of these little tubes, however, were to be found

only on one flanking branch of a rhabdome belonging to var. acadica,

which closely resembled Wiman’s Dictyonema rarum** (pl. 12, fig. 10).

I was able to trace back this composite structure up to the second theca,

but it was by no means recognizable either on the sicula or on any adhe-

sive organ. That, furthermore, the wrinkling of the surface has nothing

to do with any stress, is proved by the fact that a true cleavage some-

times does exist, cutting the widely scattered rhadomes on a slab in one

and only one direction. It produces a very fine cross striation which, in

the beginning, I was inclined to regard as a true ornamentation; but

when R. Ruedemann kindly called my attention to the possibility of the

other explanation, a renewed study soon brought me to accept it.

Another old observation, that below the fan-like tissue of a Dictyo-

nema, the counter wall of the funnel can be obtained by careful prepara-

tion, which I always found true among my material, was elucidated in

another way, because I got several specimens which had been quickly

buried and compressed in such a manner that the apertural spines of the

thecee of one wall make their appearance in regular, pointed rows between

the meshes of the covering side of the funnel. Very often, too, the

_ variety ruedemanni has its rhabdomes so badly pressed together that it

20. FRECH and F. RopmMer, Lethea geognostica, part 1, p. 570. 1880-97.

*1 CARL WIMAN: “Ueber die Graptoliten,’ Bull. Geol. Inst. Univ. Ups., Vol. 2, pl. 12,

fig. 10. 1895.

148 ANNALS NEW YORK ACADEMY OF SCIENCES

becomes rather difficult to keep those of the two sides distinct, on ac-

count of the shape and density of the branches.

Mopbe or Lire or DICTYONEMA FLABELLIFORME

Writers differ greatly in their opinion regarding the mode of life of

Dictyonema flabelliforme, one considering it planctonic, another ben-

thonic, but in recent years the idea of an epiplanctonic” life, 7. e., by

adhesion to floating seaweed, has been introduced with success by Wal-

ther and Lapworth. Even, however, in the case of Dictyonema, where

among the Dendroidea the epiplanctonic theory might possibly have

applied, an earnest objection was raised by Wiman when he showed that

D. cavernosum had branching stolons, partly strengthened by radial ribs,

leading him, as it did Jaekel before him, to the conclusion that this spe-

cies, like the “sessile denizens of the deeper regions,’ must have formed

meadows on the bottom. Matthew found at Navy Island two mature

forms, each with a distinct rootlet, and yet he made the suggestion that

“Mossibly these processes may have had some other office than that of

anchoring.” Ruedemann, agreeing with him, lays special stress on the

fact that all well-preserved specimens from Schaghticoke, whenever re-

taining more than the sicula, were provided with long thin nemas. Thus

he is inclined to assume a suspended life for Dictyonema flabelliforme,

with fixation in old stages, as unimportant exceptions. Recently, Wester-

gard, to whom we owe the best work on the Swedish material, completely

disregards the indications of what was called a “rootlet,’ stating that

among one hundred specimens no nema and no disk has been observed

in spite of distinct proximal parts exhibiting free sicule.

With these contradictory facts in mind, when studying the Columbia

collection, which fortunately preserves the two known examples of root-

lets, I proceeded to investigate all the available material with the fol-

lowing results: Of 500 specimens part way through the first stages of

growth, only 20 per cent have a well-preserved sicula, which even in

ephibastic stages ends, as a rule, in a sharp point; a few specimens, most

of which are not quite ephebastic, are provided with a nema as in Ruede-

mann’s examples and I observed only three nemas up to 20 mm. in

length; the younger the stages, the more individuals possess nemas;

78 per cent have a more or less broken or concealed end, about 2 per cent

other basal organs.

In order to present a fair account of the last, I have tried to draw all

the examples in question with most careful accuracy. (Plate X XI.)

2 Walther used the term pseudoplancton; the term epiplancton has been introduced

by Grabau to cover organisms living on or attached to pseudoplancton.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 149

In this connection several observations seem noteworthy.

1. Number of basal organs: var. conferta 2, var. ruedemanm and

ruedemanni m. f. acadica 6, acadica juv. 2.

2. At least one neanastic specimen of var. ruedemanm, 10 mm. long,

shows clearly a perfect stipe with a fringed, attaching expansion at the

base.

3. Nema and stolon-like stem have the same structure. They grad-

ually increase from .1 to .6 mm. in width and are traversed by a central

canal.

4. Disks may be expanded from the nema, the stolon-like stem or

immediately from the sicula.

5. When basal stipes are present, the sicule are rarely kept distinct.

6. Budding from stolons is not an unlikely occurrence.

A few detailed observations may give confirmatory evidence. As to

point 6, a young individual of var. conferta (11 mm. in length) has

attached to the top of its sicula a small body .5 mm. in length, from

which an irregularly fringed stem rises in the opposite direction, while

to the left denticulated processes depart. I consider it quite possible

that these denticles mark newly appearing thece which budded from the

stolon-like stipe, as in the case of Wiman’s Dictyonema cavernosum ;

but the unfavorable kind of preservation as well as the singularity of the

‘specimen fail to furnish sufficient certainty of this explanation (see plate

ROME 1/5),

Figs. 3b and 4d on plate XXI are the types of Matthew (1895) which

belong to the variety rwedemanni and are crowded on the same surface

with 16 other Dictyonemas. Only four of them have distinct proximal

parts, one a simple sicula, one (see fig. 4a) exhibits a relatively thick

stem distally broken off, the other “rootlets,” the distal portions of which

are not quite evident. Sicule are lacking, apparently from being over-

grown by the mass of the stem.

The individuals of figs. 3a, 4b can be explained as provided with true

floating organs. The specimen of figure 4¢ is excellently preserved, there-

fore of particular importance.

Now, Ruedemann, though fully discussing the great probability or, as

I think, necessity of the suggestion of a true sessile life in the later

Dendroidea, objects ((1), p. 579) that “no cases of actual attachment

and fixation have yet been recorded.” Hence, I searched among modern

benthonic forms provided with similar organs for attachment and _ be-

lieved I had succeeded, when I visited the excellent collection of the

Smithsonian Institution at Washington. On the Gorgonias there ex-

hibited, exactly the same kind of stems, partly stout and short, partly

150 ANNALS NEW YORK ACADEMY OF SCIENOES

elongated and relatively thin are observable and they rise from the same

terminal expansions characterized by irregular shape, fringed margins,

corroborating ribs and rootlike filaments, all that as changeable as re-

quired by the nature of the point of fixation. JI mention, especially,

Paramuricea borealis (from 200 fathoms) and Acanthogorgia armata

(from 160 fathoms), from the Gloucester fisheries. The last, a large,

dendroid form, is furnished with a basal stem extending distally into a

rather thin, irregularly shaped expansion which adheres by means of

two dependent lobes to the rounded edge of a bowlder about four inches

long. The similarity of this attaching instrument to that of the Dictyo-

nema figured plate XXI, 3b, is of great significance. According to my

opinion, the only reason that we have not yet succeeded in finding actual

fixation of the Dendroidea lies in the fact that the sediments of those

places on which the dendroids actually flourished were not favorable for

preserving the delicate graptolites, while in the common “graptolite

shales” we do not have the sediments on which the Dendroidea actually

erew, an assumption strengthened by the mode of embedment discussed

in the next chapter.

To get the significance of the foregoing statements, I may explain

them in the following manner: Of all distinct varieties from Navy

Island, those which tend to a callograptoid and dendrograptoid aspect

also present the greater part of adhesive organs; these are not confined

to the gerontastic stage and not separable from the nemas; the great

variation of basal organs imply a similar variation of habitat.

Here, too, we must bear in mind the results of the last fifty years”

work, as follows:

Supposed Cambrian ancestors of Dictyonema, with sicula, planctonie.

Harliest Ordovician, Dictyonema flabelliforme; young stages with long flexible

nema and thin adhesive disk, ephebastic stages generally with free sicula,

one variety sometimes sessil and that possibly beginning in neanastie stage.

Later Beekmantown, Callograpti®; free siculze sometimes observed, particu-

larly in young stages, later robust stems or hydrorhiza commonly occur.

Later Ordovician and Silurian, Dictyonemas; no nema and almost no trace of

a free sicula observed; on D. cavernosum branching stolons with buds;

on the whole, 12 species** are known provided with organs adapted for

constant fixation. Odontocaulis ; robust main stem, from which new thecze

arise. On almost all other genera of the Dendroidea, especially Dendro-

graptus no free sicula known, an abundance of organs of attachment de-

seribed.

*3 Compare C. salteri, grabaui, elegans, radicans.

ba ee a : :

Dictyonema areyi, bohemicum, cavernosum, crassibasale, desmoidea, leroyense, paral-

lelum, percrassum, polymorphum, subretiforme, stenactinotum, tenellum.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 151

I cannot help feeling that all these facts urge the conclusion that in

the history of one large part of the ancient hydroids, there was one route

of evolution prescribed, namely: The change of habitat starting from a

holoplanctonic and passing through a partly epiplanctonic to a definitely

sessile mode of existence.

ON THE OCCURRENCE AND STRATIGRAPHICAL RANGE OF THE VARIETIES

oF DICTYONEMA FLABELLIFORME

Because, fortunately, large slabs up to a size of 2 square feet were col-

lected, I was enabled to attend to the association of the Dictyonema

faunula. This is imbedded in an extremely homogeneous, highly car-

bonaceous black shale of finest grain. Small concretions of iron pyrites

are frequently scattered over the surface, though infiltration of the fossil

material is rarely to be found. The more profusely the animal mass lies

buried in the slates, the thinner they split. Cleavage of any considerable

amount rarely appears, though often recognizable in minute pseudo-

striation running in the same direction over all rhabdomes of a slab.

The slates are exactly of the same character as those of the Diplograptus

geminus—zone (Llandeilo) from Fogelsang (Skane, Loc. E5 of No.

40 of the Guide of the International Geol. Congress, 1910), those of the

Upper Graptolite-shale (Tarannon) from Stommen (Westergoetland)

or those of the upper Hartfell-shale (Caradoc) from the Moffat district

(zone No. 8, South Scotland), except that most of the European occur-

rences have been a little more affected by stress.

Two different kinds of embedment can be noticed. Slabs on the sur-

face of which single specimens with remarkably fine preservation are

found, accompanied by clusters of early stages, alternate with others

which are completely covered with a pell-mell of fragments and indi-

viduals, often almost indeterminable and lacking any favored direction.

Here the variety ruwedemanni and Staurograptus are extremely common.

On a single slab 1.5 feet square, I numbered more than one hundred

neanastic and ephebastic dendromes, omitting the rhabdomes pieced. As

a rule, both sides of the colony are closely pressed together without any

interbedded layer of mud, a strong evidence of rapid sedimentation,

caused by rough and sudden events. On the contrary, the slabs of the

first-described mode frequently show a rather thick sheet of mud inserted

between the walls of the dendrom. Sometimes this is imbedded at a

distinct angle of up to 5 degrees, found between the axis of the colony

and the surface of the split shale which is likely to be caused by the

weighing down of the heavier, distal end of the body. A rather slow

deposition has to be assumed in these cases.

152 ANNALS NEW YORK ACADEMY OF SCIENCES

As a matter of special interest, but also of insufficient evidence, the

strange arrangement of several specimens around a common center

demands careful attention. It was originally figured by Salter and

Goeppert. Afterward, however, it was questioned by the majority of

authors, but was recently again mentioned by Fearnsides ((3), p. 307)

and I likewise have observed some examples, particularly of early stages

which can be classed herewith. Fig. la on plate XXI presents two

Dictyonemas opposite each other, each of them attached to a minute

common disk. Fig. 2¢ shows two young individuals joined to each other

by means of their proximal parts. I observed also four full-grown speci-

mens of the variety acadica m. f. ruedemanni, uniting as appears in the

text-figure 2. In all the examples studied, no nema and only once a

common disk is present, so that an imme-

diate fixation by the basal parts of the colo-

nies must be assumed. If this is the fact,

which I am rather inclined to hold on ac-

count of the specimen discussed, a colonial

arrangement of the third order possibly ex-

isted comparable with that,of Diplograptus

pristis.

The statements thus far made about the

association of the faunula with Dictyonema

Fic. 2.—Dictyonema flabelliforme flabelliforme do not vary to any consider-

var. acadica m. f. ruedemanni able extent, provided that we will except

ve the fossils gathered from interbedded limy

nodules (“orstensbollarne”). G. F. Matthew°® pointed out that in the

Dictyonema-shales of New Brunswick, two bands (b, ¢) are distinguish-

able, namely, a lower one with var. conferta as the predominant type

besides var. acadica and a higher one containing an abundance of the

var. acadica and also var. norwegica. At Navy Island, both zones fur-

nish the “Bryograpti” and a few brachiopods, the latter being better

represented on the section of McLeod brook. In the slate belt of Hast-

ern New York, Ruedemann,”* though not able to fix the relative position,

again recognized the Dictyonema bed separated from the Stawrograptus

bed, which also contains Dictyonema flabelliforme var. acadica, together

with several species of brachiopods, identical with those of Matthew.

As to the European occurrence, the best recent explorations were made

by Moberg (2) and Westergard (4), who generally agree with each other

23 See references (10), (11), (12) and “On a New Horizon in the St. John Group,”

Can. Rec. Science, Vol. 4, No. 7, p. 339. 1891.

* Cf. footnote No. 15.

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 153

in keeping distinct the lower level of Dictyonema flabelliforme forma

typica which is accompanied by Obolus salteri and appolinis and a large

Lingula ? corrugata, and the higher zone of Bryograptus kjerulfi,

which has associated with it Clonograptus tenellus, Dictyonema nor-

wegicum and Lingulella cf. lepis. It is Westergard who interpolates

between them a separate zone of Clonograptus tenellus and Bryograptus

hunnebergensis. My own observations harmonize with these statements

in a quite satisfactory manner, for I found that occasionally individuals

of all described species and varieties were met with on the same slab;

that, however, in one part of the collection studied, Dictyonema flabelli-

forme var. acadica and var. conferta prevailed, associated with Monobo-

lina refulgens, while, in another part, Dictyonema flabelliforme var.

ruedemannt was mixed with var. acadica, norwegica, desmograptoidea

and Lingulella ? cf. lepis, but that in both cases Stawrograptus occurs

frequently. Although, of course, I am not able to make any statements

about the relative age of these two beds, the close conformity of the

facies on both sides of the Atlantic, when considered in a general man-

ner, seems to me to urge the assumption of homotaxial relationship,

provided we are willing to hold to the slight faunal differences due to

the geographical separation, for the supposed evolution of the Dictyo-

nema follows exactly in the same order as the historical succession.

Hence, the identification of Matthew’s zones with the two lower beds of

Westergard seems, according to my opinion, to be correct.

SIGNIFICANCE OF THE VARIETAL RANGE AND ITS PHYLOGENETIC VALUE

Before balancing the observations on the Navy Island faunula, some.

general questions bearing on the zodlogical point of view must be dis-

cussed. The first one has to deal with the term “variety,” applied to the

observed changes of Dictyonema flabelliforme. The following points

come into consideration: The features in change are not only confined

to one or the other character of Dictyonema flabelliforme, but, as shown

by the diagram and textfigure 3, every feature of specific, even of generic

value is subjected to a more or less extended change. While a small

part of the observed varieties are mostly restricted to later stages of

Dictyonema flabelliforme, in some cases, e. g., var. ruedemanni, and

desmograptoidea, a considerable telescoping of the varietal features ap-

pears, which thus have already become inherent. Furthermore, not all

of these varietal features are interchangeable; for instance, links between

var. ruedemanm and var. norwegica never occur. On those links, how-

ever, that are found, even new crossing features are observable (pyriform

154 ANNALS NEW YORK ACADEMY OF SCIENCES

shape of var. ruedemanni m. f. conferta). That this variability is of

rather important value is illustrated by the fact that the same features

which characterize the varieties are to be met again, somewhat further

developed and more distinctly separated, in later genera, e- g., Callo-

graptus, Dendrograptus, so that these features have true orthogenetic

significance.**

Considering only this point of view, we could come to the conclusion,

as did Moberg in 1906, that these modifications are useful for specific

separation. We must bear in mind, however, that all the varieties start

from a common center, 1. e., var. acadica = forma typica of the Huropean

localities; that, therefore, an unbroken chain of bridging types exists

which embrace all the different forms which elsewhere, when isolated,

were scarcely regarded as being in any way related one to another.

fof

agri

Dictyonema

Callogragtas

Tesmograptus are

rectilingatum

murrayt

2 y

D. Ss)

EE } 23

>@S. Ge =

Ma Za ce nee

oD a

Fig. 3.—Range of variability of Dictyonema fiabelliforme

Finally, any separation by time is significant only between the yar. con-

ferta and most of the other varieties, which all have apparently grown

together. For the separation of types, I regard the foregoing as of

varietal, but not of specific or mutational value. For this last, we may

regard either the saltative character or the interval of time as the essen-

tial point.

As to the cause of those changes, we may find it by starting from a

mechanical point of view. How do sessile organisms react to a more or

less continuous pull? If the pull is small, there results elongation of

27 The constantly increasing number of Dendroidea, found in the Ordovician of North

‘America, aS compared with their sporadic appearance in Huropean localities, makes it

probable that the continuous evolution was more likely confined to the American seas.

: Ii y

j gy MMI

Characters

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 155

the organs combined with elasticity and flexibility; if the pull is too

great, only concentrated forms may exist. Near the boundary line of the

trees of Alpine regions, we find species with slender, movable branches

(Betula, Larix), while in places relatively protected from storms, we

find dwarf trees and bushes, low and closely clinging (Pinus, Picea) or

extremely thickened stems on the slopes of hills exposed to windy squalls.

Water-plants lengthen their stalks or build a concentrated mass accord-

ing to their standing places. And the same reaction must be true in

benthonic animal life. The whole dendrome of Dictyonema was sub-

jected to the pull in water, 7. e., the mechanical stress exerted against

the fixed body by the action of currents, waves and breakers and thus

the animal mass proves to be influenced.

As a result of this we have:

I. Var. acadica. This middle type of all the varieties shows very

great progress over the small bryograptoid ancestors of Dictyonema, as

is proved by the neanastic and nepiastic stages, for the elastic cross-

threads enable single branches to grow to such an extent that, for ex-

ample, a colony 160 mm. long could have 20,000 theca and yet have

sufficient firmness for existence.

The varieties given here show the complete possibility of improve-

ment, but that acadica has not reached the highest place is proved by

the fact that of all perfectly preserved examples of later ephebastic

stages, acadica has less than 10 per cent, the pyriform paratype up to

50 per cent, ruedemanni up to 50 per cent, conferta 70-80 per cent,

norwegica less than 10 per cent, desmograptoidea less than 10 per cent.

Forms like acadica take the middle line of all true Dictyonemas

through the Silurian, Devonian into the Carboniferous (compare reti-

forme, scalariforme, stenactinotum, spenceri, leroyense, brairt).

II. Var. norwegica endeavors to progress by a heavy thickening of

the rhabdome net, but it causes great weighting down of the body and

much profusion of organic matter, so that this mode of building up will

be possible only when food is abundant. Compare D. quadrangulare,

murrayi, crassum, arayi.

Result: A few sporadic species up to Devonian time; no generic evolu-

tion; one-half of one per cent of our forms, more or less broken in

pieces.8

IIL. Var. desmograptoidea has closer and even elastic branches join-

ing with fewer cross-threads.

28 Also Westergard mentions the scarcity of unbroken specimens of this variety (4),

p. 60.

156 ANNALS NEW YORK ACADEMY OF SOIENCES

Result: Through forms like irregulare, homphrayi, generic evolution

into Desmograptus up to Devonian; one-half per cent of our forms,

because of somewhat later starting of evolution.

Good stolon in later species, sessile; finally, thinning of rhabdomes

(tenuiramosus, a sessile pseudo-Dendrograptus).

IV. Var. rwedemanni. Heavy branching and a successive diminution

of the elastic cross-threads produce a thick brush and dense crowding of

nourishing thecz, while still allowing movement of the rhabdomes.

Result: 35 per cent of whole fauna; 50 per cent perfectly preserved.

Half of the observed attaching organs belong to this variety. Hvolu-

tion: (a) persistence of type in a few Ordovician and Silurian species;

stolons and roots common. Compare delicatulum, pereaile, rectilimea-

tum, subretiforme, tenellum, filiramum. (6) Further reduction of

cross-threads, enforced reduction in size. Intermediate forms: Callo-

graptus, Rhizograptus, Odontocaulis, Callyptograptus; sessile, in early

stages vagrant; then Dendrograptus; stolons and roots. Further evolu-

tion of Dendrograptus leads to thin flexuose types (1.- e., under slight

pull) or to succulent types (7. e., under heavy pull) ; latter forms some-

times with central axis. End of this evolution: Ptilograptus, Acantho-

graptus, ete.

V. Var. conferta, cylindrical growth and close network, giving me-

chanically the best type.

Result: On one side 6 per cent of our forms and 70-80 per cent per-

fectly preserved; on the other no specific evolution at all, probably on

account of very unfavorable conditions of food supply.

VI. Last type of change by thickening and dividing of the cross-

threads (groups of peltatum, cervicorne, cavernosum, tuberosum) not

represented in our fauna. Appears first in American Ordovician. Sto-

lons and roots well developed.

Considering this mechanical starting point, it becomes of interest that

the Gorgonias of equivalent habitat (disregarding the funnel shape of

the Dictyonema colony) present the same lines of changes as those

sketched in the foregoing for Dictyonema flabelliforme. Thus we find

among the Leptogorgias, L. eximia and media with a network like that of

var. acadica, Leptogorgia agassizi with close fine meshes comparable to

those of var. conferta, while Leptogorgia rigida looks like a callograptoid

type and Hugorgia multifida like a Dendrograptus with beginning central

axis. The latter, still further developed, gives Pterogorgia acerosa,

while Gorgonia flabellum has a typical desmograptoid appearance and

Gorgonia quercifolia resembles closely the norwegica-murrayi line. I

cannot help thinking such a conformity indicative of parallelism in

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, VN. B. 157

evolution and that it represents one of the usual lines of development of

such benthonic forms under a given environment.

Now, with all the foregoing statements, we may touch upon the final

question of the phylogeny of the dendroids as indicated by the range of

variation in the earliest Dictyonema. A certain branch of the planctonic

ancestors common to both Graptolites and Dendroidea was evidently

pushed forward in the lines of directed evolution by the formation of

dissepiments as a supporting mechanism of the elongating rhabdomes.

This represents the Dictyonema stage, beyond which a group of retarded

species and genera (e. g., Desmograptus) never passed to any consider-

able extent. While in the Dictyonemas of the early days, fixation was

realized only by means of a thin, fragile nema, the adhesive organs were

now brought into vital and ever increasing significance. For gradually

thickened stems with basal expansions, with stolonial ramifications, with

ability of independent budding of thece and of colonies, were built,

while the original planctonic period of life becomes shortened to its final

disappearance (Dendrograptus-stage). From this point on, an extreme

widening of the main stipe, on one side, gave rise to the Galeograptus,

Discograptus, Cyclograptus, Rodanograptus-group, while a thickening

of the central axis led to forms lke Inocaulis, Acanthograptus, Cacto-

graptus, Paledictyota, to which even Chaunograptus, Corynoides and

Thamnograptus may be related, as held by Ruedemann. Finally, in types

lke Mastigograptus, a striking approach to the present hydroids has been

revealed. When considering the various races of Dendroidea on such a

broad basis, the various genera do not of course mean anything else than

stages in development; and every line of separation seems an arbitrary

one, aS 1s shown in a comparison of the species thus far assigned to

“one genus” by the different authors (e. g., the Callograpti and Dendro-

graptv). Nor is the difference between the forms in the early Ordovician

with free sicule and those with unknown sicule a reliable one upon

which alone to base the natural classification.

Within the last five years, two papers of such importance regarding

the differentiation and evolution of the Dictyonemas have been published

that they must be considered with great care. In 1907, W. S. Fearnside

(8) made some striking suggestions. He believes that

“in the earliest Dictyonema, the cells are very indistinct and rarely project

more than about a quarter of the diameter of the common canal; cross-threads

thin and numerous; stipes close together, parallel, branching at all levels;

elongated rectangles of meshes. This supposed Dictyonema diverges in two

distinct families, one approaching the true graptolites (Dictyograptus), the

other seems more nearly related to the Dendroids (Dictyonema, sensu stricto).

158 ANNALS NEW YORK ACADEMY OF SCIENCES

Dictyograptus Dictyonema

“Cells: Well-marked; tend to become “Cells: Small, generally disposed at

uniserial. angles of about 120°. Crinkly lon-

gitudinal ornament appears.

“Shape: Like a fisherman’s net from “Shape: Basketlike, starting from a

a sicula of no great length with long narrow tube or nema and di-

primary branches, diverging at an verging at angles which in the later

angle rarely greater than 90°. forms approach 160°.

“Dissepiments: Their development “Dissepiments: Their importance, in-

ever more and more delayed until crease and the general aspect of

they become practically abortive. the later forms is that of a square

or rhomboidal mesh in which cross

bars and stipes are of approxi-

mately equal importance.

“Type: var. acadica forma typica.” “Type: var. norwegica.”

There is one point in which I completely agree with this author, viz:

the importance of the differentiation among Dictyonema flabelliforme.

We both believe in this as the point of generic divergence; but in detail,

I cannot help stating that all my observations run along the opposite

direction. I merely recur to the facts that

Var. conferta, restricted to a lower horizon, according to the sugges-

tions of Matthew, Fearnside and the majority of the Scandinavian

authors, shows clearly the tubulose structure and mode of growth and

dissepiments of “Dictyonema” and the projecting cells of “Dictyo-

graptus.”

Var. desmograptoidea has cells, mode of growth like “Dictyonema,”

cross-threads like “Dictyograptus.”

Var. acadica has cells, sicula, like “Dictyograptus,” tubulose structure

hike “Dictyonema.”

Var. norwegica has cells like “Dict, POTS dissepiments like

“Dictyonema.”

Var. ruedemanni has sicula, Cements, structure ike “Dictyo-

graptus,’ cells and growth like “Dictyonema.”

Hence, as all the known varieties are in part distinguished by the

features of “Dictyonema’ and in part by those of “Dictyograptus,” such

a separation, of course, seems impossible. Furthermore, the ancestors

were doubtless bryograptoid with very distinct cells, without numerous.

dissepiments. Nor did Westergard agree with Fearnside’s assumption,,

so that against this hypothesis there appears to be no further objection

necessary.

On the other hand, Westergird, from whose detailed work my own

observations do not differ widely, starts with the following suggestions =

HAHN, DICTYONEMA-FAUNA OF NAVY ISLAND, N. B. 159

Among a hundred examples of Dictyonema flabelliforme examined, he

failed to observe nemas or disks or any adhesive organs in spite of dis-

tinct proximal parts with free sicule. He regards the wrinkling of the

rhabdomes as produced merely by pressure, and the early appearing

divisions of the nepiastic thece as due to rapid budding of nourishing

theeew. He holds it probable that the early Dendroidea like Dictyonema

flabelliforme when derived from graptolites must have possessed thece

hike those of Bryograptus kjerulf. From this point on, he considers

that there existed a great difference between the Dictyonema flabelli-

forme and the “Dictyonemas” of the later Ordovician and Silurian, as

described partly from sections by Wiman and others, and he introduces

“Dictyodendron” for the later forms.

The following objections, however, must be made after careful con-

sideration: 1. Young individuals of Callograptus (salteri in Ruede-

mann’s monograph, grabaui as described in this paper) exhibit distinct

sicule and even nemas. 2. True adhesive expansions occur among the

Dictyonemas of Navy Island. 3. The tubulose structure is by no means

referable to-any kind of stress, because this, when exerted, produces a

fine striation, running in parallel fashion over all specimens of one

slab, but cutting, of course, the tubes at quite different angles. The

associated Stawrograptus, moreover, never show a similar structure. |

4. The young stages of Dictyonema jlabelliforme, as fully discussed in

the foregoing, are found to be clearly distinct from all stages of Bryo-

graptus, as they exhibit a more primitive character (dependent growth)

than Bryograptus (declined or horizontal growth of the first branch-

lets). Hence the ancestors of Dictyonema flabelliforme were not at all

true Bryograpti, but simpler types with features bridging over those of

Dictyonema and Bryograptus.

Thus I feel quite certain that the progress of evolution which Wester-

gard believes to have existed between the early Dictyonema flabelliforme

and the so-called “Dictyodendron,’ did really take place among the

varietal series of the Dictyonema flabelliforme.

Finally, there is one point left which thus far I have deliberately

disregarded in order to simplify its consideration, for it adds merely

some complications without modifying the preceding conclusion. There

is no doubt of the fact that most of the dendroid genera have the charac-

ters not only of stadial, but also of collective groups. Of this James

Hall was partly convinced even in the ’60’s, but it has only been recently

fully substantiated by the masterly works of Wiman and Ruedemann.

Among the Dictyonemas, one part has thece with sharply prominent

160 ANNALS NEW YORK ACADEMY OF SCIENCES

lips, another possesses obscure pits and grooves in place of thecal aper-

tures; one group is distinguished by an uniserial, the other by a biserial

arrangement of the thece; a few species belonging to widely separated

divisions are proved to be heterothecal, while others are strongly sus-

pected to be homeceothecal.

We find similar differences among the genera of Callogeapinel ap-

pearance, particularly among the Dendrograpti, the Ptilograpti and in

the Galeo-Disco-Cyclograptus group. Those differences are not confined

to a certain geological age. Early as well as later species may show the

same kind and arrangement of the thece, while congeners reveal the

difference as great as possible in this respect, e. g., the highly variegated

Niagaran dendroids of America. This difficulty of a heterogeneric but

parallel evolution seems still further increased in complexity by the

repeated development of features either retrogressive or intermingled.

For we observe the strongly differentiated genus Dendrograptus already

at the base of the Ordovician (if not earlier) and yet we see that the

path of varietal evolution is in the same direction in much later strata.

We observe true Dictyonemas, the neanastic stages of which suggest cer-

tain ancestral relationship to members of the Desmograptus or the nor-

wegica-murrayt group. That the same is true among the Callograpti is

proved by a comparison of Pocta’s, Bassler’s and Ruedemann’s species

herein referred to. Even the characteristic bifurcation of the apertural

margins and dissepiments has now been found in quite different species

and genera, e. g., Ptilograptus, a very doubtful and not unlikely retro-

gressive genus, to which some Callograpti described by Pocta bear an

undeniable similarity.

However complicated the paths of evolution of the Dendrordes may

be, paths which can be traced back only after the manner of Wiman’s

and Ruedemann’s keen-eyed and careful investigations, I am convinced

that, generally speaking, they followed directions similar to those mani-

fested pre-figuratively in a small but equivalent ratio by the range of

variation in Dictyonema flabelliforme. |

PLATH XX

RANGE OF VARIABILITY IN DICTYONEMA FLABELLIFORME AND ITS EVOLUTIONAL

SIGNIFICANCE ‘

Sa!

Varieties of Dictyonema flabelliforme and Callograptus grabani n. sp. cor-

er species and. genera sketched after Hall, Ruedemann and Wiman. bata

(Reduce all given magnifications by three-fourths. )

i ’

4 f : n

een & eae ier

SAP RRD ELOY OD CATE bh? BILBAO DASE ASE OTD BEE 1! ESE,

aie

§; >

VLAD TA SOL \

he me AMI (2 a Tato, ala VON aOT Hae AAC GW, Bats wO ys We

gui raid ridatidaw to soissllor oot ti anentisod, Leute tite

Rae? Dek stirsuipsheun Hidhtievlsdsiede esonce bak

fl.

‘aes

os as

ming |‘

’ ot

Oa A TE IO, A

rue

SRS

SHR Caw

59 ont oT Be ie

Annats N. Y. Acap. Sct. Ms | | a ——— ?

oa - x I IN ;

3 4.15 ATH Ha \ :

H 1

HAG

HUH

varconterta

x4.

Crassum

vadrangularis

o / x4

a |

—

DICTYONEMA. -——--==— a J DICTYONEMA

leroyense

| peltatum

tuberosum v

N a

x20.

—

GZ Zz =

ZZ gat

——

mt acacia --canterta

/] cermoorne 4 CQVeINOsum

\ hal

» /FLABELLIFORME |

WV aca C7 rectifineatum\\i{}}

14 Wi

DICTYONEMA

__ subretiforime

‘4

Octal

PTILOGRAPTUS

poct

X70.

XA oes ACANTHOGRAPTUS

SMO pane TUS * SUCCICUS X20.

D xzo "“f flvit tans

P \ succulentus

AF:

. \f

tenviramosus ©... gicus DENDROGRAPTUS

x4 DENDROGRAPTUS : os

PLATE XXI

BASAL ORGANS OF DICTYONEMA FLABELLIFORME

1 a, b, var. conferta

2 a, b, c, acadica (0, c, juv.):

3 a, b, acadica m. f. ruedemanni

4 a-d, ruedemanm

All specimens are preserved in the collection of Columbia University and

are enlarged five times in the figures. The measurements given refer to the

length of the entire colony, which is not always represented.

49 iu aa ae det Om Gt ak wy <i hLonneogey | sep al rate te

DRO PLO ACT ALLTOP Rs AVY ACE SE Sb Fy

VA ASS

Hii Veter stitaiow ‘te 1)! nS ih Der MeoI1c 8 i

Si) OL MRIOT toy Ringe ges: odd. aeMEgét

PHMSMOTEOT BERLE TON BEATE

WA |}

IXX Giv1d ‘TIXX @Wwa70A ‘IOS “dVOV “A (N STIVNNY

PLATE XXII °

ULAR COMPARISON OF THE SPECIES OF DICTYONEMA, DESMOGRAPTUS AND

CALLOGRAPTUS

is ae 4

HA LOGHe He Th VO PrAL Mons %

sje

BO ETTARDORIAD

We soln eae beuttoltaidwry, : ee

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gitdsngy to Taai+>

: WAL (Gate

; BREESIIOY, oiCl «

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Yiodae 2aawel & ey

ren: eros

jaa et tpadastgs

? 1 sto I ee Seu Bs i eek ret bes (8

fo. Sica sem | sdial Yo; She 8

(BLEE #,

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rsacrid -)- . brisitiosG wok oe

dalinunc’ ct

nn a ee ee

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jcmioesetea len : "Beoxiatt

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)

Annas N. Y. Acap. Sct.

ee eee cn 0

VoLtuMB XXII, Puatr XXII

Rhabdomes Dissepiments Thecee Notes

Length’ Per cent

Names Shape and size ee Fi Se | | Number Diverg. | Number Basal organs

individual a a nies

width i in Character se on . in Gharacter Meshes Number Character Passing into Char. of stage Evolution and age | Preservation

Dictyonema flabelliforme | infundibuliform and : early stage bryo- | center of variabili

ey pyriform 16:1 9-10 4 (.8-.5) mm. 45° 45 thin quadrang. 16 sicula, nema conferta penta Me GEV sites 90% of lat

60% projecting : apache true Dictyonemas e FaveRHG:

acadica large—very large 9:1 (8-12) subparallel 80° (3-8) 2mm. (elongated) | (15-17) ?1 stolon (young stage) | esmograptoidea | tate stage with F

Me ruedemannij} t stages broken

mixed features | up to lower Carbon-

| iferous

infundibuliform 8:1 4 mm. Ais 4 isp nga ,

brairi 8-10 25 4-5 .25 mm. CLES Menger le ct mocKe..! i) = onpactvoginte el -- otene all <2 Sc nesoac Species r

medium size (?) subparallel lower Carboniferous DOken

| infundibuliform 6 45° basal adhesive neanastic

leroyense 1:1 8-10 s 2-4 .25 mm. subquadrang. | ..... | 9 «ss... Fea tee a | ee eT with thickened Onondaga broker

large ? ee 80° - disk branches a Cy

vasiform—cylindri- 2.5:1 4 (.38-—.5) mm. 45° 7 thin atarie 7 delicate atari Gh early stages:

cal (3.7:1) gy, 2 quaoreng: Sua BOGIES bryograptoid; a 20% of lat

y. conferta 6% 8.5-10 BuBDAY 1 basal disk ruedemanni 2 not any evolutio % of later

parallel— = 4 1-2 mm. z . + oti : . 4 ; late ephebastic y » h. st:

medium size (large) | (1.5:1) parallel ie (8) regular (GERM) | (UGE) | apsaktegtas ?1 branching stolon | (pyriform link) contraction iis Seen

5 are .5-.7 mm. a5 thick— a ; char. of var. a few species in

ee infundibuliform re um ars 50 6 irregular rectang. re pues a nCadiCn usually restricted Ordovician; Aen

: large * robust 90° (4-8) subcircular acute (conferta) to late stages of yarallel evolution up

8 + parallel (.1)-.6 mm acadica 1 F

+1) =: p to lower Devonian

2 e 1.5 mm. 50° rigid

murrayi , 2 4 sient hae 4 7 rectang. 9-10 MCuUber we PSA TS Sree atetel ate poeeema |b eke Madsen et | Beekmantown = broken

arge ‘ .4-.9 mm.

parallel

| infundibuliform 1:1 Bb alr 50- irregular rounded distall

crassum | 9 Rone CG Yara ee ea emma aot 4 eR ol lee cee ||.8 stance lt. MIeDROUGnDDTOT ae wll ~ sabaroAe. denmig mantel New Scotland + broken

| large (?) + parallel (90°) 4-9 mm. subquadrang. rap!

cba of var. ri

.4 mm. usually restricte

infundibuliform 8 + 90 3-4 very irregular irregular delicate to late stages of to genus Desmo-

v desmograptoidea 1:1 YG subparallel— IGT] | ee oe a? adn onepoodoeds acadica acadica ; graptus, up to + broken

medium size (7-9) flexose— (45°-90°)} (1-5) .1-4 mm. subovate less projecting middle Deyonian

joining acceleration be-

ginning

| infundibuliform tent very rare elongated appressed basal parts with

y 8 F o a + well

intricatus 2 ¢ 1:1 15-20 fils (64ND) 45°-90° (0-1) tet agama aes ered S85 : siculay?’- - —mylia 3 Bares Dictyonema char- Chazy preserved

a medium size coalescent) irregular subovate circular acter

Z ae =

a f eg 8 chitinous basal

2 infundibuliform 21 ; very rare elongated appressed . =

cancellatus A 1214 | adulating (thick) | = 90° | (0-1) frequent Seo Sai Airey gale Al). uenee mbar t + broken

medium size (?) yw ; sae 5 irregular subovate circular aOTiatann

sicula, emir aiamen Gah similar species up

infundibuliform— Segall ess FI SAS ESS to Devonian

y. ruedemanni OLY (J) 33% xz Be + 45° : 7 pars | eee 15-16 delicate | Dee orstolon\withidisk, see nee! ete lati 4070 or Tess

6 = ‘ * urther coolution roken

large—very large (1.1:1) (10-15) subparallel (1-5) -lmm. elongated branching stolon in (pyriform link) | old Biased parely atin (OFiboe Func

; neanastic stage RoR BEER’ Dendrograptus

zi 2 4 irregular old stages partly

rectilineatum 2? 12-14 ai) Hel st 45° 3-5 quadrang. | ..... irregular ?shortstem | .seeee desmograptoid Chazy broken

medium size au parallel “et —.16 mm,

° eyatiform 1:1 4 45° delicate; quadrang. 13-14 3 A partly well

gracile etek 10-12 (6 at the base) 14 BSE projecting Dasalestem and dickoue | eeanmesee eee | ERE Niagaran preserved

: a J ] 700

palette (HE) (@) subparallel 10, -1 mm. elongated (207)

- =

B 3-5

. a infundibuliform irregular : partly dendro- Beekmantown— partly well

salteri 5 11 14-15 subparallel st 45° O(-1) very few off case stec 14-18 thin hydrorhiza | ...... graptoid Chazy preserved

3 small—medium size (flexose ) appressed

a Sw ls Se ei a

lisey

les

‘.

—fividl

iauibana

arotilsd ibaritess

open ete ctte tee 5

acotifdibantai

satel

arrdtified ibastat: 2

ogiel

uel she a Steers

; as]

idee ce : =

eae 1 &

ee i: | a:

! 3 =. j H Ps)

~ Pees bs 2

poet EY | |

4 H 5 — } : |

: : : } ; {

SScannES aeiaet CalsiarienaericemeaeE a rel u oe AS ee a Cl Ce PRON ew 8 TN Se tel Le ew Fa eat

| | | | | 8 ~ | | | |

roe ; | CI OO cael ; | i

Y : —~ Rio * Se —. i s

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oe o ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXII, pp. 161-224

Editor, EpmMunp Otis Hovry

METAMORPHISM OF PORTLAND CEMENT

ER ape) ae eae oe eee Be” eee

ALBERT B. PACINI

NEW YORK

PUBLISHED BY THE ACADEMY

10 SEPTEMBER, 1912

ee Se Oe Pay RED Ee PP ee

oe ae a re. . y

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Narurat History, 1817-1876)

OFFICERS, 1912

President—Eurrson McMituin, 40 Wall Street

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P bg

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ae eee he IPM lle Bee heehee Eien Steg

The sessions of the Academy are held on Monday evenings at 8:15 . :

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Natural History, 77th Street and Central Park, West.

[ANNALS N. Y. Acap. Scr., Vol. XXII, pp. 161-224.

10 September, 1912]

METAMORPHISM OF PORTLAND CEMENT?

By Apert B. Pacini

(Read before the Academy, Part I on 8 January, 1912; Part II, 1 Apri,

Introduction........ é

Nature of the problem

CREM AMCOMNMPOSMMOMerts seis. 5 slic sca es oecle cies hae ce gals se cetncan ce ee ee. 164

liltmerealoecall COMSAT 6605 6 Sabon oe Doo oe lo Oeibo- co Dom cco mn ic pico ood t5 Ios

SHAMS [PROEESS. o ceccwls 6 pe GORE CO DO De BO CIG Ore Dirks o.coie ner aeiricne eno. clarion 165

TEIAIRCWAINTING?: (AIROVGES Sc Gc Glo.c co GIES b CIn IRIS DIOIGs DIC EnDICIC ISI oi oleic CioloiCi ina ie erciicicioae 166

Influence of water upon metamorphism. ...............0.- 2 eee cece eee .. 168

Temperature of the water at first added......................02005- 169

Temperature of the water that may eubseutenuly come into contact

Brats Na TehT CRSA LTA eearee ty eters US at craia celle level e le -eversifelsigiesaiele ois ic elaeid.w ele 171

Ant VyAOkeWwAlel: ab MES Ad Med so.5% 22 cnelaels © oe cece se sacle s dec ebisces 172

SUAS Ol CAmSMe [INANE Gonobo somos odUn Oe CbOUmoSlnd oon GoIOoccs 172:

BAAN COs ees! 25, ce cis aryans Ses 204 5 SE cs ROE RE LS OE Cs ae 173

EDV CODY SISt UNC Ol yeeicna cel etetety orate ae) ooo ate ala ccy2)ci/dici ceils sta elae oda eee Cues 174

Mechanical agitation when water is added...................... 174

Total quantity of water at first added.............. Sethe eiele vs ae 175

Quantity of water that may subsequently come into ponte with the

SV SECIS ie sreter ciel eteiere a fata oes Ao Cee SHESat a CORE GOR Oe ae Ee 175

Steel COM ERE ALM CIES: sarece lateveva eretetae ale Senses oer aipieve fa Hine uvenacs sae erets T/T

Membranes........ Hed coosobodooeaubscese soc ORD ES ooage spec soaoo UIT

MAIS S eG eat Gl USivers.srttereeusirct apiece hoc tencheeanteeo fave Ua aitezeilel-cilenuieveraavere rele Sateen eli

Quahity of water at. first added. es ee dee ela a ee ah ate Tendele us excole eee 178

EwinemMmaterial: iW SOlIiONNs 932522 o us cleo oe seasons coe wees 178

Quality of water that may subsequently come into contact with the

SWSUCMM.c.0, e205 < ele 5S EOC O RO es OR A DS See rune ree ENC UEP AER USE Mr oa RA Sige OLS a 180

Senne TMA Th ODIO Goo ou doonomoc cuss cos obo db donee ous eso

SS Ge NPC Te syeira cea crete Me cape\vetna. ours nc eeSueare araiabevonehal eapeuaueaicanstaets PEO eco 180

Alailicangd! deep TOck WATS? Hiss cocis shGesiete = bes oats secceperene Sree leo

Having material in suspension

1912)

CONTENTS

eee ee eee see eee eee ese eee

1A thesis submitted in candidacy for the degree of Doctor of Science at New York

University, 1912.

Acknowledgments are due to Prof. J. Edmund Woodman and Mr. Raymond B. Earle,

of the Department of Geology, New York University, and to Engineer Inspector Ernst

Jonson, Board of Water Supply,

City of New York, for valuable suggestions made dur-

“ing the preparation of this paper; also to Mr. Fred H. Parsons, Assistant Engineer, and

Messrs. James E. Jay, Charles M. Montgomery and Charles EH. Price, Inspectors, of the

Board of Water Supply Laboratory, for material assistance during the experimental

work.

(161)

162 ANNALS NEW YORK ACADEMY OF SCIENCES

Part II

Page

Experimental investigation. 0.5. s he cs ele Oo ees See Oe 184

Temperature of the water at first added.................. ...cccceee 186

Temperature of the water that may subsequently come into contact

With: the Systems. 2). 2005.04 00s. shia cele aeete ole eee 186

High-pressure ‘steam: .. 6. 20025 Sosa a eters seins eee 186

Cold: storages io sae accisie's Rae SE Oe U CE een eee A 187

Quantity of water at first added...................--00+- ceccceccce 189

Size of cement particles... 5. 22.26.5 5. acanc uses Bae 189

Mechanical agitation when water is added...................... 191

Setting time of cement in laboratory air and in damp closet...... 192

Effect of excess of mixing water on strength of concrete......... 193

Effect of excess of mixing water on permeability of concrete. .... 194

Effect of excess of mixing water on strength of neat cement..... 195

Effect of the presence of clay and dissoived substances.......... 198

‘Quantity of water that may subsequently come into contact with the

SY SLCIIE ahaa eh ate een etc isalSila bien Seen en eee 200:

PerMe aD ity. ccsyé.ois5.0 oeananes oslo koe ioe sto wee DS ne ee 200

Coneretes containing different aggregates.....................-- 204

Concretes containing different cements...................-+eeeee 205

Effect of the direction of flow through concrete.................. 206

Quality of water at first added). . .....0.6 2. tc eee eee we eles | ole ene 208

Compressive strength of neat cements gaged with various solutions 208

Effect of gaging with various solutions upon the strength of

mortars afterward stored in water.............6....----+-ee> 210

Effect of gaging grout with rock waters.................-.---+: 211

Quality of water that may subsequently come into contact with the

SV SUC. wc. d05 aie: dite ais loves. « ecanai a wh cea eysh w re tetane ne mite tevlerele teee ee one eI I eke neem 213

Theoretical considerations. .............- 2c cee eee e ee tweet ee ees 213

Effect of storage in various saline solutions upon the strength of

GW TINOZNRS Sagoo das losSoncouseceoodcedonsobes oes es ioe ee 214

Effect of storage in rock water upon the strength of lean cement

LINOLEIC ESPRIE See eI AIas REN ones ence AIO OAc ACO O COOKS 216

Summary of experimental results................... “beets ia Cae Lee Se RS 218

General GCOMCIUISTONG!. c.5 clos voce okeies e cee cle eile aos sei) le iovedenslololejehckers i) tea Reems 219

ISN HON Nhs onldeoGakcomdoocedonGbOos Bou Guu DOOgnUe Oo DOIUc OSG Cb O00 settee 220

PART I

INTRODUCTION

The important field of investigation covering the changes which take

place in the setting and hardening of Portland cement and in Portland

cement which may be considered to have attained the greater part of its

maximum hardness calls for the services of experts in various branches

of science. Many of the general problems can, as a whole, be relegated

PACINI, METAMORPHISM OF PORTLAND CEMENT 163

to the petrologist and hydrologist; and this paper is an attempt to treat

cement as a rock, differing from other rocks only in being artificial, but

subject to the same internal and external influences as other components

of the earth’s crust.

_ A training in geophysics and geochemistry is, perhaps, the most valu-

able asset in surveying the field of Portland cement. If no other end is

achieved by the following pages, the mere representation of the question

as a problem in applied petrology will, it is hoped, help future investi-

gators in a more systematic inquiry.

Part I of this paper is devoted to a necessarily brief review of the

present status of the subject, and no attempt is made to discuss data

quantitatively. Experimental results in elaboration of the various points

discussed are presented in Part II.

The experiments described in Part II were made at the laboratory of

the New York Board of Water Supply by the writer, and in part by his

associates, in the course of the investigations of the Board. The most

modern and complete equipment was available, thanks to the prudent

foresight of the gentlemen at the head of this great engineering enter-

prise. The data are reproduced by permission from the periodical bulle-

tins of the Inspection division and from the annual report of the Board

iow alae

NATURE OF THE PROBLEM

Portland cement is a finely ground artificial rock, whose essential

constituents are silica, alumina and lime. In it are found a number of

component minerals recognizable by definite optical properties, but the

individual constitution of which is not yet clear. The percentages of

these minerals vary somewhat according to the method of manufacture

and the purity of the raw materials, but there is, on the whole, a fairly

stable proportion in a series of normal cements.

The method of manufacture of Portland cement will not be discussed

here further than to state that it consists essentially of the calcination

of a mixture of calcareous and argillaceous rocks at high temperatures.

Usually, about 2 per cent of gypsum or of plaster of Paris is afterwards

added to retard the set. By varying the proportion of these rocks, the

temperature and duration of calcination, the fineness of grinding, and

also by the addition of foreign substances products are obtained having a

wide range of hydraulic properties.

The hydraulic properties are setting and hardening. Setting is the

attainment of rigidity by the plastic mixture of cement and water and

begins immediately after mixing, requiring several hours for completion,

164 ANNALS NEW YORK ACADEMY OF SCIENCES

Hardening is the progressive increase in strength acquired by the mass,

and it attains the greater part of its ultimate value in about a year.

Even after this period it is subject to a small progressive increase (42).

In general the properties of setting cement are to be found both in

mortars, or mixtures of cement and sand, and in concretes, or mixtures

of cement, sand and broken stone, these chemically inert materials added

to the cement exerting a physical influnce on metamorphism.

CHEMICAL COMPOSITION

The chemical composition of normal Portland cement is shown in the

following tables:

Average of 300 Normal American Portland Cements, Representing 20 Brands

of All Types

(Analyses By the writer for the Board of Water Supply)

Maximum Minimum Average

Osc siases srs oho ksucavlin ce coreneaey en aolene nce ahetioe erence: 25.89 19.85 22.70

1 EH O en aceon ntens areca tees MRR Ur eA oe 4.08 1.23 2.73

NTO ee apeaie Mopaeds os tees wee atm catdene Aree, 9.16 3.43 6.17

CAO R etars aniitecets eacedercoe tay seensieeetee eae ans 64.91 59.06 62.67

MES ©) iain 2.2 cde ae ty on eaten eae 4.00 0.30 2Zald

SOs sctacercasut tater a acre taruelens iene pace 1.75 0.84 1.37

COREG O; faillcalieSs. Sc .cseacy oe cree eke cacao oie es ea ee nase Pas lerl

Average of 100 German Portland Cements

(Burehartz, (12) )

STO esas a ceie te catacter a vonblesoraniane ay teats eeebone SPO touell oceania rece eRene, Sane 20.87

1 er! 0 aaron mane rnin ace Sri er eae cinein a Sects bod Ss 2.98

U2 O eae Cees a BIER, eM onl NN Gan entire cin Same m emote’ o Oe 7.63

C621, etre Anse ernment ena Taree Mn RES frre. ce cea BRA Gc 62.99

MeO asic, At Besealereredtates nade eG) ae cere eect aarate nea ete ac ne emepesie ave 1.55

SO hace a Bhs aie a toe ocean ata seanie acy raat cet ta ores aps aaaviopesallatariey amet aeeas 1.85

The ultimate chemical composition of a cement is only, however, a

rather indirect clue to its hydraulic properties, just as the ultimate

analysis of a composite rock may only give a faint idea as to its con-

stituent minerals or possible products of metamorphism. For example,

it would be quite possible to synthesize a mixture which would, on analy-

sis, correspond exactly to the chemical composition of an excellent Port-

land cement, yet which, when gaged with water in the ordinary way,

would develop practically no tensile strength, in fact would possibly fail

to set at all.

Cement, therefore, must owe its hydraulic -jseifbill 2s to a particular

grouping of its constituent compounds, quite analogous to a series of

2 Numbers in parentheses refer to the bibliography at the end of this article.

PACINI, METAMORPHISM OF PORTLAND CEMENT 165

minerals; and looking to the identification and classification of these

minerals, a great deal of investigation has been done.

By trial burnings of simplified mixtures, such as lme-silica melts, and

by microscopical examination of sections of the resulting clinker, the

problem is gradually being clarified, but, owing to its great complexity,

much controversial literature thereon has been issued on both sides of

the Atlantic (52, 69, 80, 64, 65, 88). The theories put forth have so far

had little practical effect upon the manufacture and composition of the

commercial product (63).

No complete and final enumeration of the chemical duipotiine result-

ing from the burning of such a mixture of clay and limestone has yet

been accepted as authoritative. The microscopical identification of the

individual chemical compounds which go to make up the mineralogical

entities is at best somewhat unsatisfactory, especially because of the

minuteness of the particles of raw materials necessary to secure thorough

and uniform calcination, and consequently the extremely small size of

the resulting crystals and aggregates. It has been proposed, in this con-

nection, to secure these of a size available for study by the expedient of

fusing the clinker in an electric furnace; and, by this means, a partial

clarification of the system has been obtained (103).

MINERALOGICAL CONSTITUTION

The minerals which are recognized in cement clinker have been named

alit, belit, felit and celit (101), and a metamorphism? of these occasioned

by the action of water is the cause of the setting and hardening of Port-

land cement.

Alit has been reported a solid solution of tri-calcic silicate in tri-calcic

aluminate, and celit a solution of di-calcic aluminate in di-calcic silicate

(61). Other investigators have reported alit and celit to be silicates of

different silicic acids (26).

Belit is probably a calcium aluminum silicate of the composition

Ca,Al1,8i,0,,, a form found in nature as the mineral gehlenite (27).

SETTING PROCESS

Precisely what chemical reactions and physical transformations take

place in the setting and hardening processes is not yet definitely settled.

It may, however, be stated that by modifying the proportions of clay to

limestone through a certain range, we obtain a product which varies in

its speed of setting and of hardening. In general, cements high in silica

3 Metamorphism: Any change in the constitution of any kind of rock, Van Hise (104).

166 ANNALS NEW YORK ACADEMY OF SCIENCES

are found slow setting and slow hardening, while those high in alumina

are quick setting and quick hardening. An increase of lime in the latter

retards the setting (63).

The calcium aluminates are probably the main factors in the setting

of cement, while the hardening is due to the calcium silicates. The mag-

nesium compounds are inessential to the hydraulic processes (105).

Upon the addition of water to cement, the equilibrium in the system

of solid solutions and chemical compounds is destroyed, and a series of

changes is inaugurated tending towards the production of a system which

will be stable under the new conditions. The first effect resulting from

the solutions and reactions brought about by the presence of water is the

setting of the plastic mass.

Under ordinary conditions of practise, the quantity of water used is

about 22 per cent in the case of a neat cement, being less in the case of a

mortar, and still less in the case of a concrete. When this proportion of

water is used, it is probable that the setting of cement is mechanically

analogous to the setting of plaster of Paris and is caused by the growth

throughout the mass of a network of crystals, deposited from the satu-

rated solution formed by the first stage of hydro-metamorphism.

Owing to the low solubility in water of the original component sub-

stances, the attainment of final equilibrium is a matter of considerable

time, and is further delayed by the automatic protective action of films

of insoluble substances coating the active particles (23). 'These films in

some cases are semi-permeable, and exert a selective influence upon the

solutions osmotically penetrating them. Under normal conditions, that

is under those conditions which have been found in practise to yield the

densest and strongest product, this attainment of equilibrium considered

apart from the setting process at first proceeds rapidly, but the rate of

increase of strength grows smaller, tending to a minimum.

A. Erskine Smith has shown (90) that there has been no permanent

retrogression in the strength of cement in the case of specimens kept

under observation for 21 years. Of course, this relates to laboratory

specimens protected from weathering, but shows one of the directions

which this metamorphism may take.

HARDENING PROCESS

The hardening of cement has been ascribed variously (48)

1. To the fineness of grinding,

2. To the increasing stability of calcium compounds due to combina-

tion of part of the silicic acid as the silicates grow less basic,

3. To the action of free lime upon calcium compounds,

PACINI, METAMORPHISM OF PORTLAND CEMENT 167

4. To the decomposition of basic products present in the freshly set

cement,

5. To equilibrium of calcium hydroxide with the siliceous constitu-

ents, and

6. To the hydration of the double silicates and anhydrides of lime and

alumina.

The two theories that have at present the greatest claim upon con-

sideration are that the strength of set cement is due to the progressive

erystallization of calcium hydroxide (80), and, in some respects diamet-

rically opposed, that this strength is due to the formation of a dense

complex colloid, soft at first but gradually adsorbing calcium hydroxide

and thus becoming harder and harder (64, 65).

According to the latter theory, cement consists of a mixture of fused

compounds of silicic, aluminic and ferric acids with lime, together with

an excess of lime, partly dissolved and partly enclosed. Upon the addi-

tion of water to this system it is decomposed, and the water becomes a

supersaturated solution of salts, which react between themselves. The:

compounds resulting from these reactions crystallize about the cement:

grains in needle-shaped crystals. So far, the process is analogous to the

setting of plaster of Paris (45), and silica takes no part in these pre-

liminary reactions.

A hydrogel begins to form about each grain, in which the crystals

become embedded. This hydrogel consists essentially of calcium hydro-

silicate, and to a minor degree of calcium hydroaluminate and calcium

hydroferrite. At first it is soft and plastic, but gradually becomes dense

and rigid by the adsorption of calcium hydroxide. The strength of

cement is mainly due to this process of coagulation.

The calcium hydroxide may of course erystallize and lend additional

strength; but its crystallization is rather more hkely to burst the har-

dened cell walls about each grain of cement, and thus admit liquids

later in the process which may be fatal to the integrity of the structure,

either by undesirable chemical reactions, or simply by dissolving away

the lime, with the formation of soft hydrates of silica, alumina and iron

oxide, instead of the desired hardened colloid (64, 65). -

Much corroborative evidence has been offered by supporters of this

view, and similarly by the exponents of the crystallization theory in

defense of that. The question is still at issue, and the main difficulty is

the microscopic recognition of the constituents of set cement (34, 78).

Unquestionably, colloidal materials result from the action of water on

silicates of this type, when the particles have been ground to the fineness

of Portland cement (23, 21,95). This has been directly observed in the

168 ANNALS NEW YORK ACADEMY OF SCIENCES

case of cement and reproduced with synthetic mixtures. What binding

power colloidal material may develop is strikingly seen in the case of

conglomerates and sandstones in which hydrous silicic acid, aluminie

hydroxide or ferric hydroxide has been the cementing material, so that

the theory is attended by a high degree of probability.

On the other hand, it is also quite conceivable that the interlocking of

crystalline masses between the grains of cement may account in some

measure for the strength. There is definite evidence that calcium hy-

droxide does crystallize, and its mineralogical and crystallographic con-

stants have been determined (24).

The two views are not entirely irreconcilable, and it is possible and

even probable that, mechanically, the strength of cement acquired by

hardening is due to both processes. Whatever be the chemical reactions

in detail by which these elements of the structure are produced, the main

condition for their occurrence is the presence of water.

This paper is devoted to an enumeration of the factors which influence

the metamorphism caused by water in Portland cement, and the varia-

tions in the physical properties of the resulting rock, brought about by

varying these factors.

INFLUENCE OF WATER UPON METAMORPHISM

The action of water upon Portland cement is a resultant of

1. The temperature of the water

A. At first added

B. That may subsequently come into contact with the system

2. The quantity of water

A. At first added *

a. Size of cement particles

b. Mechanical agitation when water is added

ce. Total water added

B. That may subsequently come into contact with the system

3. The quality of water

A. At first added

a. Having material in solution

B. That may subsequently come into contact with the system

a. Having material in solution

b. Having material in suspension

* Owing to the peculiar autoprotective reaction of cement against the action of water,

before alluded to, the quantity of water coming into contact with cement is a function

of the size of the particles and of mechanical stripping of protective films, as well as of

the ratio of cement to water.

PACINI, METAMORPHISM OF PORTLAND CEMENT 169

The final effects of geological processes do not differ in the main,

whether these operate upon natural substances or: upon the products of

human industry. The agent whose activity is responsible for the majority

of terrestrial changes, namely water, is also the main factor in the meta-

morphism of the artificial rock, cement. By intelligent control of the

action of water upon this rock, the desired results are obtained, and its

value as a material of construction is inestimable. Lacking this insight,

the action of water may result in catastrophe, or at least loss of time,

money or efficiency. Geology, then, through hydrology (59), is enabled

to give substantial aid to the engineer.

TEMPERATURE OF THE WATER AT FIRST ADDED

In construction, the water at first added to cement, known as the

gaging or mixing water, is subject to the entire range of variation of

atmospheric temperature. The lower limit is far below the freezing

temperature of water and of course, in this phase, water is useless for

the purpose.

Within the possible range of temperature under working conditions, it

has been established that as the temperature of the gaging water used is

higher, the set becomes more rapid. Considering the setting due to the

deposition of a network of crystals from the supersaturated mixing

water, the beginning of this deposition would be sooner attained, if the

water reached its condition of supersaturation more quickly; and this

condition would be brought about by a higher original temperature,

provided, of course, that the solutes increased in solubility with the tem-

perature. With a higher temperature, the volume of the water would be

greater and the viscosity less, and consequently its range of activity would

be increased ; that is, it Would be enabled to reach a larger number of

cement particles and thereby more quickly arrive at its saturation point,

and the deposition of the crystalline network hastened in consequence.

If the temperature of the mixing water be above about 37° C., the setting,

instead of being hastened, begins to be delayed. If the deposition of this

network were a simple case of precipitation from a hot solution, it would

be logical to state that the solubility of the compounds concerned was so

high at this temperature that they were not deposited from solution.

The problem, however, seems chemical rather than physical, and it is

more probable that this effect is due to hydrolysis.

Hydrolysis increases with the temperature. In the case of the weak

salts that must exist in the system we have under consideration, the ulti-

mate products of hydrolysis are the gelatinous materials—silica, in the

hydrated form, aluminic hydroxide and ferric hydroxide. The adsorp-

170 ANNALS NEW YORK ACADEMY OF SCIENCES

tive and coagulative properties of these materials unquestionably do not

compare with the coagulative powers of the complex colloid which

Michaelis postulates (64, 65). If, therefore, the hydration of cement

does not proceed in a properly regulated manner, it is conceivable that it

may become a hydrolysis, with deleterious effects.

If the mixed cement is allowed to freeze, the setting will not take

place, but on thawing out the mass, setting is resumed. Obviously the

transition of the water to the solid phase hinders solution and diffusion,

and upon resuming the liquid form, water promotes these processes as

before. A slow setting has, however, been observed in frozen mixes (94),

and it is quite possible that the phenomenon of regelation may account

for this.

Smoke gases have been found to have a disintegrating effect upon

cement setting at a temperature lower than 7° C.; this is attributed to

the formation at these temperatures of a hydrated calcium carbonate,

having the formula CaCO,,5H,O by the action of the carbon dioxide of

the smoke gases upon the lime of the cement. At slightly higher tem-

peratures this hydrate is transformed to pulverulent calcium carbonate,

with consequent disintegration of the structure of which it forms a part

(107).

The effects of moderate variations in the temperature of the mixing

water upon ultimate strength are practically of no great moment; even

mixes that have been frozen and afterwards allowed to resume their set

are not materially affected in their ultimate strength, if the set has not

proceeded too far at the time of freezing (11). More than one repetition

of the freezing process upon the same mix, however, will be quite de-

structive to the final hardening.

If the hardening be considered a process of. crystallization, repeated

freezing may be assumed to destroy the strength by the formation,

through rapid temperature changes, of relatively small and non-adhesive

crystals of the calcium hydroxide during the critical foundation period

of growth of the crystalline structure, so impeding and misdirecting

consequent interlocking that a weak structure results.

If, on the other hand, the colloidal theory is adhered to, it is only

necessary to point out that the colloidal cell walls about the cement

grains may be ruptured by the expansion of the contained water in freez-

ing. This would result in discontinuity of the internal structure, and if

sufficiently widespread, as would be the case in repeated freezings, would

alone account for weakness.

Studies have been made of the ultimate resistance obtained from frozen

mortars by varying the amount of gaging water, with the view of estab-

PAOCINI, METAMORPHISM OF PORTLAND CEMENT 171

lishing whether “wet” or “dry” mixes best resist the disruptive effects of

frost during setting. The results reported are discordant. An excess of

water has been found by one investigator to enhance the effects of frost

(85), while by another it has been found to diminish them (11). Theo-

retically, the disruptive effects of freezing should be enhanced by the

presence in the mass of larger quantities of gaging water. On the other

hand, it can be assumed from the colloidal standpoint that an increase in

the amount of water present will result in the formation of a greater

quantity of colloids and a greater elasticity of the resulting mass, to-

gether with a smaller total breakage of cell-wall material.

TEMPERATURE OF THE WATER THAT MAY SUBSEQUENTLY COME INTO

CONTACT WITH THE SYSTEM

The action of hot and boiling water upon set cement is strongly

marked in the case of cement which contains free lime, producing after

a few hours, swelling, distortion and cracking and even total disintegra-

tion. A normal cement so treated, however, preserves its original form

and volume after short periods of exposure to the boiling temperature.

The viscosity of water at high temperatures is greatly diminished, and

the liquid is thereby enabled to penetrate more rapidly the capillary and

subeapillary voids, thus reaching more quickly a larger internal area.

If, as in the case of an unsound cement, free lime is thereby reached, this

is slaked much sooner than it would be under normal conditions, and

moreover with great violence, owing to the higher temperature of the

water, producing internal disruption, and perhaps thus opening up fur-

ther avenues to the penetration of water, with a repetition of the slaking

process.

The boiling test here described is a very important one in the testing

of cement for construction, but it is perhaps less reliable in the case of

unsoundness from the presence of excess of magnesia.

In cements stored in waters of relatively high temperature, it is prob-

able that the processes of solution act more rapidly, from the two reasons

mentioned above; but evidence is lacking to show that any significant

decrease in ultimate strength is thereby occasioned.

Data as to the storing of cement in waters of low temperature, yet not

subjected to the action of frost, are not available in the literature, but

they would be interesting.

In the case of exposure to the action of frost, the process is quite

similar to that which goes on in the disintegration of natural rocks and

depends, in like manner, upon the initial mechanical resistance of the

172 ANNALS NEW YORK ACADEMY OF SCIENCES

mass, upon the total volume of the voids and upon the ratio of capillary

to subcapillary voids. The disruptive effect is, of course, due to the ex-

pansion of the water during freezing. Consequently there is a possibility

that during the earlier stages of the history of the mass this effect may

be to a great extent neutralized by the presence of soft colloidal material

(45), because of its lack of rigidity.

Voids are undoubtedly present even in neat cement mixes, and they

are more common in mortars and in concretes; when, therefore, these

have attained a sufficient hardness, they are in all respects similar to a

natural rock and subject to the same katamorphic processes. The effect

of frost increases in intensity as the mass ages and loses elasticity.

As water permeates the cement, even after hardening has progressed

to a considerable extent, it becomes charged with various electrolytes, and

its freezing point is consequently lowered. ‘To some extent this immu-

nizes the mass from frost action. On the other hand, as we have seen

before, cryohydric compounds may be formed at these low temperatures,

and the separation of these from solution is a factor in the opposite

direction.

QUANTITY OF WATER AT FIRST ADDED

Size of cement particles—tThe finest particles in cement, provided that

they are chemically identical with the remainder, are the most active

cementitiously, because of the ease of reaction and of the greater proba-

bility of this action being uniform throughout the mass of each particle.

‘This is recognized under the microscope by the ultimate disappearance

of these particles as individuals upon the addition of water. Owing to

the relative insolubility of the constituents of cement, both before and

after metamorphism, each particle becomes covered to a certain depth

with the reaction products, which in this case take the shape of gelatinous

films (2) in such manner as to offer hindrance to the further action of

“water.

The particles whose diameter is smaller than or equal to the thickness

of this zone evidently are the most efficient chemically. The larger par-

ticles are less so, as the passage of water through the enveloping film is

a slow matter, and some particles may be so large as to remain internally

unchanged. It is probably this fact that gives a hydraulic quality to

previously set cement that has been reground and retempered with

water ; in fact, this process may be repeated a number of times with the

same sample of cement.

Not all of each particle, therefore, can take part in the setting and

hardening, and sometimes this proportion of inert material is consider-

PACINI, METAMORPHISM OF PORTLAND CEMENT 173

able (86). The coarser particles are comparatively inert and might be

replaced by grains of foreign material of the same size without ma-

terially influencing the ultimate strength of the resulting mass. This

has been demonstrated experimentally (17). It does not follow, how-

ever, that a cement consisting entirely of uniformly very fine particles

would be a desideratum, since such a cement would not pack as well as

one containing a greater variety of sizes, and the increase in chemical

activity would be markedly overbalanced by the imperfection of struc-

ture of the mass. Considering each particle to be spherical, and of equal

size with every other, when packed in the most compact manner possible

the pore space would be nearly 26 per cent (89). The points of contact

of the adjacent spheres, notwithstanding the tendency of the gelatinous

envelope to spread, would be relatively few. If, however, this pore space

were filled with finer material, the structure would develop more strength,

The function of part of the cement is to remain passive and to add to the

strength of the structure merely by its action of void-filling. Extremely

fine grinding has been found to decrease the ultimate strength, if the

cement is used neat, but to give greater strength, if the cement is used

in a sand mortar (62).

As might be expected from the above considerations, the fineness of

erinding has an accelerating effect upon setting. Cement ground in a

tube mill until only 1 per cent remained on a sieve having 5000 meshes

per sq. cm., was so quick setting that it could not be restrained even by

the addition of 10 per cent of gypsum (47). When cement is relatively

coarsely ground, the ultimate strength is not so quickly attained, but its

acquisition is regular and uniform!

Laitance.—In concrete construction under water, especially salt water,

there gathers about the freshly deposited concrete a milky white cloud of

suspended matter, technically known as laitance. This material is also

formed when concrete is mixed very wet, though not deposited under

water.

An analysis of laitance by the writer, made for the Board of Water

Supply, practically coincides with an analysis made by Richardson (97)

and leads to the same conclusion as that reached by him; namely, that

laitance represents an actual loss of cement and consists of the finest par-

ticles of cement which have been washed out of the concrete. The addi-

tional conclusion is justified that this portion of the cement, by reason of

the small size of its units, has been so acted upon by an excess of water

that it has undergone complete hydrolytic decomposition, before the col-

loidal enveloping film had adsorbed sufficient electrolytes to completely

coagulate it and so render it largely impermeable. This is substantiated

by the fact that laitance possesses neither setting nor hardening qualities.

174 ANNALS NEW YORK ACADEMY OF SCIENCES

Hydrolysis theory.—The formation of such a protective film upon the

surface of a coarse particle will so regulate the access of water to its

interior that the contents will be slowly and normally hydrated. If the

entire mass of the particle were at once accessible to an excess of water,

the weakly acid and basic compounds at first formed would soon be hy-

drolysed and shorn of their binding power, and instead of the normal

complex colloids described by Michaelis (64, 65), capable of adsorbing

electrolytes and so coagulating into a dense rigid mass, simpler colloids

such as hydrous silicic acid and aluminic hydroxide would form, which

have not these powers to so high a degree.

Finally, the rate of setting and hardening of a cement may be con-

sidered a function of the proportion of fine particles present. Mortars

set and harden more slowly than neat cement, and concretes more slowly

than either. This is simply a development of the fact that coarsely

ground cement sets and hardens more slowly than that which is finely

ground. It may be considered, from another viewpoint, that the inactive

material interferes with the liberation of heat from the system, and that

chemical reaction is consequently delayed in proportion to the amount

of inert material present.

Mechanical agitation when water is added.—lf cement in the state of

a plastic mass be worked and kneaded, the ultimate strength will benefit

thereby, up to a maximum time of working. It is legitimate, a priori, to

surmise that the setting is hastened, within limits, although no record of

this is found.

After the maximum time referred to, which in experiments made at

the Board of Water Supply laboratory has been found to correspond

roughly with the time of initial set, continued working will cause a fall-

ing off in the strength. Up to this time, mechanical agitation with the

proper amount of gaging water will cause an inerease in the ultimate

strength.

The formation of the crystalline network, which constitutes the setting

of cement, and which is responsible for the primary strength by holding

the plastic mass rigid and in place, while the more important elements

of hardening make their appearance, is unquestionably facilitated by

agitation. Stirring is a means of hastening chemical reactions by bring-

ing the agents into more intimate contact. The compounds that go to

make up this network, being sooner brought into solution, perform their

function more quickly, and the crystals begin to form. Instead, however,

of forming a continuous rigid network, the crystals will be smaller and

less cohesive than if undisturbed in their growth, and the set can be

delayed and even prevented by continuing the agitation long enough.

PACINI, METAMORPHISM OF PORTLAND CEMENT 175

The ultimate resistance of cement which has been thus treated is

decreased as well. The formation of the coagulated colloid, or of the

interlocking crystal units, whichever may be the cause of hardening, is

rendered imperfect and discontinuous, and the structure reflects the

weakness of its component units.

It may moreover be supposed that more cement has been brought

within the range of hydrolysis by this agitation, and so converted into -

laitance, even the larger particles being stripped of their protecting films

by the attrition, Tests made at the Watertown Arsenal (36) showed that

after one hour’s working, cement had gained 4 per cent over the normal

strength, but that after 10 hours’ working, it had lost 24 per cent from

the normal, in 20 hours 38 per cent, in 50 hours 56 per cent and in 100

hours 69 per cent.

Total quantity of water at first added.—Under certain conditions, the

entire range of particles of a cement might be destructively hydrolysed,

resulting in what is termed “drowned” cement.’ The effect of an increase

in the quantity of mixing water is known to result in a diminution of

strength, and, bearing in mind what has been previously said regarding

hydrolysis, the reason is clear. If, before the cementing of contiguous

particles, an excessive amount of water is admitted to contact with the

cement, colloidal material will form in increased amount: It has been

shown that an increased amount of mixing water results in an increased

volume of the paste produced (39). This indicates that a larger amount

of the products of hydrolysis is formed.

Owing to difference in composition between these hydrogels and those

formed under normal conditions, they are incapable, as has been before

observed, of adsorbing electrolytes in such degree as to attain to the

density and rigidity of the latter, Admitting, on the other hand, that

colloids so formed do not differ in composition from those formed in the

normal hardening of cement, there still remains the abnormality of the

structure formed in this way. Being discontinuous, it would not offer

the same total resistance, in the form of connected films, to the passage

of water. Moreover, in the presence of an excess of water the working

ratio of electrolytes to colloids would be less because of the greater dilu-

tion in proportion to the volume of colloid.

QUANTITY OF WATER THAT MAY SUBSEQUENTLY COME INTO CONTACT

WITH THE SYSTEM

The effect of water upon cement after it has completely set rapidly

diminishes to a negligible quantity at ordinary temperatures, if the water

is reasonably free from dissolved or suspended impurities. There is a

176 ANNALS NEW YORK ACADEMY OF SCIENCES

leaching out of calcium hydroxide from the mass of the cement; but this

diminishes as the mass grows more and more impermeable, by the coagu-

lation of the colloidal cell walls and by the carbonation or other precipi-

tation of lime salts in the pores.

This deposition of lime salts in the pores is evidently the cause of

higher strength in specimens which are allowed to dry out a few hours

before testing. It is analogous to the higher strength developed by sea-

soned stone than by freshly quarried stone, occasioned by the evaporation

of the “quarry sap.” In addition, the carbon dioxide conveyed to the

material in a gaseous form is absorbed by the hme and may be considered

a positive factor towards strength, while that conveyed in solution (where

the cement is under water) is a negative factor, in that it accelerates the

solvent effect of the water coming into contact with the cement. On the

other hand, cement specimens which are entirely air-hardened are un-

questionably weaker, by reason of the absence through evaporation of the

requisite amount of water for proper hydration.

When the action of water upon set cement is intermittent, the solvent

effect manifests itself by unsightly incrustations and discolorations (3),

caused by dissolved material brought to the surface through capillary

action and there deposited by evaporation. When the mass is perma-

nently under water, these salts are merely washed away. The danger

from these incrustations, although slight, is the disintegrating effect pro-

duced by their increase in volume, through crystallization or efflorescence,

and the consequent disruption of the denser surface skin, rendering

easier the action of frost upon the entire mass.

This surface skin is improved by troweling the semen while in a

plastic state, and consists of a closely packed layer of fine particles, which

offers high resistance to permeation by water and comparative immunity

from the solvent action favored by a rough, porous or fractured surface.

If the mass be placed in water before setting, it is more lable to hy-

drolysis, as evidenced by the copious formation of laitance; and if greatly

exposed, as by agitation under water, it may fail to develop the greater

portion of its normal ultimate strength. To prevent this, care is taken,

in laying conerete under water, so to convey it that it offers the least

possible surface to water action during its descent; and to this end it is.

either lowered in cloth bags, or filled in through a chute, so as to escape

all avoidable exposure to hydrolysis.

If the water which comes in contact with a cement structure be under

considerable pressure, so that its tendency is to percolate through the

mass, the solvent effects will of course be magnified, proportionally to the

porosity of the mix; and experiments made by the Board of Water Supply

PACINI, METAMORPHISM OF PORTLAND CEMENT 177

have shown that concrete subjected to such percolation has been shorn of

the major portion of its ultimate strength. In this case, the solvent

effect of the water is only part of the influence at work, purely me-

chanical factors entering largely into the destructive process, as will be

shown later.

Stalactitic growths of lime salts form as the result of water percolating

through concrete. Micro-organisms of the algal type frequently lodge in

the pores of concrete and by their growth may act as a protective influ-

ence against the permeation of water. The effect of their products of

metabolism and decay upon the concrete structure has not been studied.

Numerous waterproofing materials and processes have been devised

(40, 73). They may be grouped conveniently under three heads.

Surface treatments.—The application to the surface of concrete of a

coating similar to a paint has the disadvantage that concrete is not a

thoroughly dry material. Where the vehicle is a liquid immiscible with

water, the paint will not therefore come into contact with the concrete

proper. If the vehicle is miscible with water, unless insoluble products

are at once formed by reaction with the constituents of cement, the

active agent is quickly leached out.

Membranes.—These are layers of waterproof tissue interposed between

two layers of the concrete. There is strong probability that these never

actually form a bond with the concrete, and thus they necessarily intro-

duce an element of weakness and heterogeneity.

Mass treatments——The active material is incorporated with the con-

crete at the time of mixing, either by dissolving or suspending in the

gaging water, or by intimately mixing with the cement or sand. These

treatments are many and differ widely in the agents employed. Sub-

stances of a waxy or fatty nature, triturated to a great fineness, are the

most generally offered, but the incorporation of these in a mass of con-

crete is generally followed by weakness of the structure. The general

problem of cement waterproofing has been conceded to be simply a ques-

tion of void-filling, yet this must be accomplished without the addition

of inert material that will weaken the resulting structure.

The addition of more colloidal material has been suggested. This is

ingeniously effected in a recent process by the use of hydrolysed cement,

obtained by treating cement with an excess of water (99). The paste so

obtained is added to the cement during mixing.

The still unclarified state of our knowledge of the chemistry of the

setting and hardening of cement is the great handicap which has thus far

prevented the devising of a satisfactory waterproofing agent. A large

number of the waterproofing preparations on the market are therefore

178 ANNALS NEW YORK ACADEMY OF SCIENCES

purely empirical, and not applicable to the practical waterproofing of

large masses of constantly wet concrete. In the interests of efficiency, it

is probably more economical to expend money destined for waterproof-

ing in the purchase of additional cement to be used in making a richer

concrete.

QUALITY OF WATER AT FIRST ADDED

Having material in solution.—On adding water to cement, heat is

evolved, the temperature of the mix rising in some cases to above the

boiling point of water. It is the custom to look with suspicion upon

cements in which an excessive rise of temperature is obtained, as being

liable to develop unsoundness. The abnormal rise is attributed in some

instances to the presence of free lime, in others to an insufficient propor-

tion of lime. The volume changes caused by a rise in temperature have

‘been given as the reason of the difficulty encountered in joining fresh

‘cement surfaces to old, causing weakness at the plane of juncture, the

‘contraction of the mass on cooling breaking the joint before it has devel-

oped sufficient strength to resist the strain.

To prevent this, it has been suggested to coat the surface to which

fresh cement is to be appled with a retempered mortar; that is, with a

cement which has been treated with water after partial setting. This

provides an intermediate course of material in which the temperature

changes are not so rapid, and upon this course the fresh cement mixture

is applied (35).

Upon the same principle may be explained the use, for a fresh course

of cement which is to be joined to some which has previously set, of

mixing water in which a quantity of cement has been stirred, thus retard-

ing the chemical reaction and consequent temperature changes. In both

cases, the active water is already charged with the soluble portion of

cement, its solvent power for the same material is thereby diminished

and the chemical action moderated, so that heat is more gradually

evolved and violent expansions and contractions avoided.

The influence of dissolved electrolytes in mixing water has received

much careful study. Through the addition of a small percentage of

some soluble salt to the mixing water, many have tried to influence the

properties of the completed structure and to produce a mass that would

develop greater strength or a higher degree of imperviousness. Unfor-

tunately, the panacea has not as yet been discovered that is suitable for

practical application.

The addition, similarly, of a soluble powder incorporated in the mass

of the cement comes under the same category. In this connection, our

PACINI, METAMORPHISHM OF PORTLAND CEMENT 179

attention is drawn to the effect of the usual addition of ground gypsum

or of plaster of Paris to the ground clinker, for the purpose of retarding

the set. There are other salts whose retarding influence on the set of

ground clinker is comparable and probably superior to that of gypsum,

but their use is not so practical, consequently, it has been adopted as the

restrainer for general use.

Tt has been shown by Rohland (83) that the salts which respectively

accelerate and retard the setting of cement are the same as those which

accelerate and retard the hydration of quicklime. From this it is con-

eluded that their influence is “catalytic.”

A detailed explanation of the mechanism of the action of gypsum has

been put forth (79), holding that the presence of calcium ions in the

mixing water, resulting from the solution of gypsum therein, decreases

the solution of other calcium ions, thus retarding the solution of hme

and the hydrolysis of the aluminates, which in turn retards the set.

It seems probable, upon this basis, that the presence of certain elec-

trolytes in the mixing water acts upon the set by influencing the solu-

bility of calcium sulphate therein, and consequently increasing or dimin-

ishing the number of calcium ions present in the mixing water as a result

of the solution of calcium sulphate.

For example, sea water has been found to retard the set of cement

(83). Gypsum, although a relatively insoluble salt, may be regarded as

fairly soluble in moderately strong solutions of sodium chloride or of

other salts having no common ion (14). In the presence of sodium

chloride, then, the: calcium ion concentration in the mixing water is

raised, and the solution of the calcium aluminates diminished, with the

effect of retarding the set. Sulphates have been found, when dissolved

in the mixing water, to have the property of retarding the set, with the

exception of aluminum sulphate and calcium sulphate when in low con-

centration. In view of the latter fact, it is evident that the above expla-

nation is perhaps only a partial one.

A large number of other electrolytes and miscellaneous compounds

have been investigated and the results are recorded (83).

The effect of soluble constituents in the sand used for making concrete

is by no means negligible (4) and may offer an explanation for many

instances of puzzling behavior of the mixture.

Sea water has been and is, in many instances, still used for mixing con-

crete, and to the best of our knowledge, no cases of failure can be attrib-

uted to this cause alone. Apart from the influence upon setting, the

presence of dissolved electrolytes in the mixing water seems to increase

the strength of cement in the early periods, as far as reported results have

180 ANNALS NEW YORK ACADEMY OF SCIENCES

shown (4). This may perhaps be due to an increase of coagulation of

the colloidal constituents, by reason of the presence of salts of greater

ionization than are generally present. On the basis of the crystallization

theory, this phenomenon is rather difficult to interpret.

QUALITY OF WATER THAT MAY SUBSEQUENTLY COME INTO CONTACT

WITH THE SYSTEM

Having material in solution.—A large number of failures in concrete

structures have been attributed to the disintegrating action thereon of

water impregnated with various salts. Inasmuch as all ground water is

charged to some degree with salts which it has accumulated in its passage

through the soil and rocks, this problem is worthy of the most careful

attention. For our purpose, such mineral-laden waters may be divided

into

1. Sea water

2. Alkali water (from western alkali soils)

3. Deep rock waters.

The mineral constituents are common in all these cases, and vary only

in the prominence of one or more of them. ‘Thus in sea water the chlo-

rides of sodium and magnesium, in alkali water the alkaline carbonates,

and in deep rock water the chlorides of calcium and magnesium and the

sulphate of magnesium are the distinctive constituents. Whether the

effect of these electrolytes is cumulative, so that the continued action of

solutions of low concentrations will work harm, or if not, what are the

limiting concentrations to assure safety to the structure, has not been

worked out. Obviously, it is not a laboratory problem, since the factors

which obtain in nature are impossible to duplicate on a small scale. The

solution lies in careful inquiry into the mechanism of the action and in

observation of the instances of failure in construction work, with a study

of its causes.

Sea water.—The effects of sea water upon set cement have been sum-

marized in the statement by Feret, “No cement has yet been found which

presents absolute security against the decomposing action of sea water”

(97). Le Chatelier, after a series of experiments extending over ten

years, confirms this conclusion (53). Poulsen concludes, however, that

the chemical action of salt water is not alone sufficient to cause Portland

cement mortars to deteriorate (76).

The diversity of results reported in the observation of the action of

sea water upon cement indicates that there are varying factors at work

that so far have not been clearly recognized. Whether the precise nature

PAOINI, METAMORPHISM OF PORTLAND CEMENT 181

of the action is physical or chemical is not quite settled. There are not

lacking investigators who assert that the destructive action is mostly

physical and is due, among other causes, to intermittent submergence

and consequent deposition, by evaporation of crystals in the pores of the

structure, which, either by their pressure of formation or by expansion

during efflorescence, have a disruptive effect similar to that of frost (98).

There are those who hold that the action is entirely physical, and is

due to this factor and the effects of frost (91, 102), although probably

the latter is seldom the case in sea water, owing to its low freezing point

{50). The effect of direct sunshine has been found deleterious when

alternating with that of tidal action (20). Undoubtedly, all of these

factors contribute to the total effect, and there is as well a marked

chemical action.

The chemical effects of sea water upon cemient are capable of various

interpretations. They are summarized as the formation of complexes by

the action of the dissolved sulphates and chlorides in the water upon the

calcium silicates and aluminates of the cement (74). It has been stated

that sodium chloride solutions have the power of dissolving calcium sili-

cate with the formation of an unknown salt (58, 70), and also that the

sodium chloride enters into combination in the mass, the chlorine ion

entering into the combination calcium chloro-aluminate, and the sodium

ion combining with lime, silica and alumina, to form compounds of the

nature of the zeolites.

Working with strong solutions of the individual salts of sea water, it

has been found that the chief harmful constituent is magnesium sulphate,

and it has been suggested that this salt reacts with the lime of the cement

to form calcium sulphate and magnesium hydroxide. The calcium sul-

phate further reacts with calcium aluminate to form a calcium sulpho-

aluminate, which by swelling causes the disruption of the mass. The

magnesium hydroxide formed has been regarded as a restraining agent,

by virtue of its filling up the pores of the cement and preventing further

ingress of sea water (70). Again, the disruption has been directly at-

tributed to the increase of volume caused by the formation of this mag-

nesium hydroxide (46). It has been calculated that, apart from the

formation of hypothetical sulpho-aluminates, a molecularly equivalent

amount of calcium sulphate replacing the calcium hydroxide of the ce-

ment occupies 2.08 times as much space and is, therefore, the cause of

the disintegration (13).

Alkali and deep rock waters——Burke and Pinckney (13) have formu-

lated a working theory of the action of the various salts common to all

natural waters, They attribute the disruptive action to more rapid re-

182 ANNALS NEW YORK ACADEMY OF SCIENCES

moval of the calcium hydroxide, and in some cases to its replacement by

material occupying greater volume, as before shown, and consequent

disintegration of the structure.

That some such reactions occur is indubitable, and that the mechanical

factors are a large influence in the disintegration is equally certain. An

additional cause which may be of great importance has hitherto been

neglected. The electrolytes in these natural waters may act as acceler-

ators of hydrolysis, and, in effect, cement which is in contact with sea

water is subject to the same action as that of an excess of water from any

cause. By the presence of these electrolytes the hydrolysis of a larger

proportion of the cement is effected; and the results are increase in the

volume of the hydrolysed portion, and production of a larger proportion

of inert colloids. It has been found that a larger amount of cement can-

be converted into colloidal matter by the presence of an electrolyte in

the water with which it is treated (99), and also that the speed of hydra-

tion of cement is affected by the presence and proportion of electrolytes

present (84). The fact that a larger amount of laitance appears to be

formed in sea-water construction also seems to bear out this theory.

Besides the reactions mentioned, set cement is subject to the replace-

ment of silicic acid by carbonic acid, as are the natural rocks. Especially

is: this true in cases where the cement comes into contact with marsh and

peaty waters and waters containing ferrous carbonate, which by transfor-

mation to the hydroxide liberates carbon dioxide (24), which has been

found to act, not only upon the calcium hydroxide but also upon the

silicates and aluminates (28).

The presence of free acids in water which acts upon the cement is

quite destructive, in proportion to the concentration of the acid and to

its strength or weakness as an acid. It is quite probable, however, that

the liberation of colloidal silica by the action of acids would serve to a

great extent as a protective influence against their further action.

Sewage gases are generally effective by reason of the hydrogen sulphide

which they contain. This gas is readily oxidized to sulphuric acid, and

then its action is the production of soluble calcium and aluminum sul-

phates, which are subsequently leached out from the mass. This action

has been found greatest at the surface of the liquid (106). Hydrogen

sulphide may also act by converting the iron of the cement into sulphide,

and this becomes oxidized into ferrous sulphate and is leached out, or by

its expansion causes disruption (28).

The action of many other inorganic and organic solutions has been

observed, but they do not come within the scope of this paper, since they

are not met with in natural processes.

PACINI, METAMORPHISM OF PORTLAND CEMENT 183

In general, the consideration is worthy of attention whether concrete

structures which are under stress are not more lable to chemical disin-

tegration than those which are in repose, or whether a single structure is

not more liable to this action in its strained parts than in those not so

affected. We have data to show that strained iron is more liable to corro-

sion than unstrained, and it has been asserted that strained minerals are

more acted upon by underground solutions (104).

A number of protective measures against the action of saline waters

upon concrete have been suggested and tried, but none has been so strik-

ingly effective as to achieve universal recognition. The simplest remedy

suggested is to make the concrete for such uses denser and more imper-

vious by the employment of a greater proportion of cement, yet this may

not always be practicable. When concrete is exposed to the gases result-

ing from the decomposition of sewage, it is suggested that even such a

proceeding may be of no avail (29).

Previous air-hardening of the concrete before laying under sea water

is acquiring more widespread use and is highly recommended (87).

The cause of its protective action is attributed to the carbonation of the

calcium hydroxide (48).

Variations in the fineness of grinding and in the chemical composition

of the cement used in concrete for sea-water construction have been pro--

posed. The French specifications for sea-water cements call for a finer

grinding than that which is required for ordinary construction. Much

has been claimed regarding the resistance to disintegration offered by the

so-called “iron ore” cement, which contains a minimum of alumina, this

being almost entirely replaced by iron.

Having material in suspension.—The peculiar nature of the series of

compounds forming and formed from cement, in that they are all of

- relatively low solubility, tends, as has been before observed, to retard the

reactions which may occur. Mechanical agitation, by promoting diffu-

sion and by transporting the reacting materials to their possible spheres

of action, will accelerate these reactions. The motion of water, per se,

can and does produce this effect, and when the water is armed with sus-

pended material, its activity in this direction is greatly enhanced.

Where water has immediate access only to the outer surface of a mass

of set cement and its pressure is low, the effect is a slow corrasion of the

dense surface skin and ultimate removal thereof, rendering the interior

gradually more accessible. Ordinarily, this process is a slow one, al-

though under certain conditions, as in coast protection works where the

velocity of the water is high and the suspended material coarse and

plentiful, the destructive effects are more to be reckoned with.

184 ANNALS NEW YORK ACADEMY OF SCIENCES

The effects from less spectacular processes are quite surprising. Where

the pressure of the water is such that there is a marked motion of the

water within the pores of the concrete, the erosion is internal and far

more insidious. In this case, the suspended material is part of the struc-

ture itself. Small particles of cement or, in the case of mortar, grains

of sand which become detached from the parent mass are whirled around

by the water stream and shortly enlarge the cavity in which they are

rotating, until it merges with some adjacent cavity. Under favorable

conditions this process may continue until the interior of the structure

is greatly weakened.

A factor which to some extent neutralizes the flow of water through

concrete is the choking of the pores by sediment, coming from the water

itself or furnished by the action of the water upon the concrete. If the

flow is oscillatory, as in concrete exposed to the range of the tides, this

protective effect will of course not be so marked (54).

Diatoms and other microscopic marine organisms with siliceous or cal-

careous tests undoubtedly play an extensive part in the preliminary-

stages of this internal mechanical action, by choking the capillary spaces.

At the same time, undoubtedly, the organic debris thus introduced may

by its decomposition give rise to substances, carbon dioxide and hydrogen

sulphide, for example, which have an accelerating action upon the proc-

esses of solution, and the silting effect may thus be neutralized or even

overbalanced.

PART II

EXPERIMENTAL INVESTIGATION

In Part I, the ways in which water may influence the metamorphism

of Portland cement were discussed qualitatively, and their possible effects

upon the permanence of the structure of which cement forms the basis

were pointed out. This question has now assumed economic and vital

importance. eS

In the following pages experimental data are offered, in elaboration of

the outline laid down in the first portion of the paper. Points in the

scheme which have been established beyond doubt by previous investi-

gators are here omitted, and only such results are inserted as have been

deemed necessary as additional evidence. The last division of the out-

line, treating of the action of suspended material in water in effecting

the erosion of concrete, has not been experimented upon, not having come

within the scope of the writer’s activities, and therefore is omitted.

PACINI, METAMORPHISM OF PORTLAND CEMENT 185

Other divisions have already been so thoroughly covered by previous in-

vestigators that very little remains to be said about them. Emphasis has

therefore been laid in this paper upon the little known fields.

The problems which confront the user of concrete are of a high order

of complexity. The generalizations of chemistry are not yet sufficiently

developed to apply rigidly to systems of so many variables, and experi-

mental work on a laboratory scale often fails almost entirely to reproduce

the conditions of practice. The best guide to the truth, then, is the prag-

matic sanction of experience—the investigator in this field can but point

out probable directions for future experimentation. The theories which

underlie past success are a safe guide, nevertheless, to future construc-

tion, and the systematization thereof is a legitimate field of usefulness.

While, strictly speaking, any aggregation of chemical compounds

might be considered a rock, whether natural or artificial, a majority of

the cases conceivable under such a classification would not present im-

portant petrological problems in the study of their metamorphism. Such

a problem as the action of water upon a mixture of sodium chloride and

calcium sulphate can be partly solved in vitro, even though the action

of sea water upon gypsum deposits is an interesting petrological investi-

gation.

The important components of Portland cement are everywhere about

us in nature, and the reactions by which it is made artificially have been

taking place for many geological ages without the intervention of man.

Silica, alumina and lime are among the most important constituents of

the earth’s crust; they are subjected in places to the same conditions

that exist in the kiln, and are afterwards acted upon by water, under

some of the same conditions under which man builds massive structures.

The complex question of the history of rock magmas is not one to be

solved by any one group of scientists, but by patient and concerted efforts

of the chemist, the physicist and, above all, the petrologist. So the prob-

lem of the constitution of Portland cement may be as yet somewhat inde-

terminate; but an examination of the more general effects of metamor-

phism may reveal some identity with conditions in natural rocks already

studied and may direct us to the correct methods for investigation of the

constitution of cement (67).

Other important problems in the field of cement and concrete are re-

ferred to in the following pages, and belong in great measure to the field

of petrology. Not the least important of these is the suitability of vari-

ous types of rocks for use as aggregates in concrete, and this work is

claiming more widespread attention daily (19, 44, 111).

ANNALS NEW YORK ACADEMY OF SCIENCES

(op)

TEMPERATURE OF THE WATER AT FIRST ADDED

Two standard cements were gaged with the requisite quantity of mix-

ing water for each at different temperatures. The effect upon the time

of initial and final set was noted, as follows:

TABLE 1

Effect of Temperature of Gaging Water on Time of Initial and Final Set

Per cent by weight Temperature of Initial set, Final set,

of mixing water mixing water hours hours

A B A and B A B AM | B

29 21 70° F. 4.2% | 4.50 | 627) |leueae

22 2] 100° F. 1.50 4.00 4.00 7.00

22 Pall 150° F. 0.33 3.75 0.50 5.75

22 21 Alm e 1.00 2.79 2.75 | 6.00

The results seem to indicate that interference of hydrolytic decompo-

sition with the setting appears between 150° F. and the boiling point of

water. Below these limits, the effect of increase of temperature of the

- mixing water, as is well known, is to increase the speed of setting (31).

The setting time at these temperatures is a resultant of two opposed

processes,—the formation of the water crystalline network, and the de-

structive hydrolytic action of water upon the original constituents of the

cement, resulting in a product which has no hydraulic qualities.

Where the second process overbalances the first is the point at which

the speed of setting ceases to increase and begins to diminish.

This is true of course of the stage known technically as the final set

(9). In the first few hours of setting, there is a period of relaxation,

which McKenna has aptly termed reverse set, and which he has been

able to detect with precision by means of an ingenious chronographie

apparatus of his invention (60). ‘The phenomenon has been observed by

the writer and his associates in the laboratory of the Board of Water Sup-

ply, using the Vicat needle; but this apparatus does not lend itself to a

scientific study of the finer differences in rigidity which occur during the

setting period. McKenna’s apparatus should throw a great deal of light

upon the initial metamorphism of cement. ;

TEMPERATURE OF THE WATER THAT MAY SUBSEQUENTLY COME INTO

CONTACT WITH THE SYSTEM

High pressure steam—Wig (109) has recently presented an account

of the excellent effects of high pressure steam when used in curing con-

PACINI, METAMORPHISM OF PORTLAND CEMENT 187

crete. He found that by using concrete that had attained its initial set

and exposing it to steam at 80 pounds pressure the six months’ strength

could be obtained in two days, a tremendous accelerating of the harden-

ing process.

This state of affairs is not very satisfactorily explained, if the harden-

ing of cement is supposed to be due to the progressive crystallization of

calcium hydroxide, since it is somewhat at variance with our knowledge

of the conditions of crystallization to assert that continuous exposure to

a high temperature, presumably constant, should accelerate crystalliza-

tion ; particularly since in this case the amount of water present in the

system remains the same. On the basis of the colloid theory, however, it

is simply explained by supposing that adsorption of calcium hydroxide

by the complex hydrogel is accelerated by higher temperatures.

Cold storage.—A series of tests, embracing neat cements and mortars,

was made upon tensile test specimens exposed, after the age of 24 hours,

to low temperatures under diverse conditions. The following conditions

were observed :

1. Chilling the briquettes at 24 hours’ age by filling the storage tank

with water at the lowest winter temperature as it came from the tap.

The water was then allowed to come slowly to normal winter temperature

for the tank, about 60° F.

2. Chilling another set of specimens, otherwise normally treated, by

filling the tank with cold water as before, 24 hours before breaking.

3. Storing another set in ice water for the entire period after remov-

ing from the damp closet at 24 hours’ age.

4. Normal treatment.

Two brands of well-known cement of high quality were run in parallel.

The mortars were of proportions 1:3, Ottawa sand being used. The

results obtained are summarized below:

188 ANNALS NEW YORK ACADEMY OF SCIENCES

TABLE 2

Effect of Cold Storage on Strength

ngt ounds

% ature of Storage “per sauuare meen ree ae Number of

Cement Mix Blorsee PaothiGd Species

deg. F. 7 days | 28 days | 7 days | 28 days

x Neat 60 Normal 730 739 0 0 10-12

43. 1 606 693 18 6 12-12

42 4 654 739 11 0 12-12

60 Normal 702 745 0 0 24-24

34 3 638 669 9 11 24-23

xX 23} 60 Normal 300 361 0 0 11-12

43 1 281 361 6 0 11-11

42 2 283 371 6 +3 11-11

60 Normal 313 408 0 0 24-24

34 3 262 312 16 24 22-24

nya Neat 60 Normal 628 843 0 0 12-12

43 ] 650 770 +3 9 12-12

44 2 649 872 43} +4 12-12

60 Normal. 628 697 0 0 24-24

34 3 530 622 16 11 22-24

Y 1:3 60 Normal 250 300 0 0 12-12

43 1 253 370 +1 +6" 12-11

44 2 287 327 +15 +7 12-12

60 Normal 228 317 ) 0 22-24

34 3 197 234 14 26 22-24

From these results, it is safe to conclude that, aside from the effects of

frost, low temperatures are adverse to the development of the hardening

process in cement, and that in general this effect is more pronounced in

mortars than in neat cement.

The adsorption of calcium hydroxide by the complex hydrogel may

proceed at a lower rate at lower temperatures; or if this is not so, the

primary hydration, of which this hydrogel is the product, may proceed

more slowly, and thus less of the hydrogel be produced,—either of which

processes will detract from the hydraulic activities of the mass. It would »

seem from the experiments that the latter is the more satisfactory expla-

nation, since the test specimens which were chilled at first and allowed

to return to normal temperature show a tendency to return to normal

strength at the longer periods, while the general tendency in the series

kept constantly in cold water is to fall further off from the normal, indi-

cating only a limited available amount of hydrogel to undergo the coagu-

jating process.

PACINI, METAMORPHISM OF PORTLAND CEMENT 189

The effect of sudden chilling at a period when a large proportion of the

strength is already developed does not show any decided direction, both

the positive and negative variations from the normal averaging the same.

It may therefore be concluded that, for the temperatures studied, a chill-

ing of this kind has no significant effect.

An explanation according to the crystallization theory of hardening

would fail to fit the facts so satisfactorily. In the specimens that were

chilled at first and allowed to return to normal temperature, there should

be under this hypothesis a more significant decrease of strength, owing

to the formation of small, non-cohesive crystals from the rapid tempera-

ture change. The return to normal conditions should not favor so nearly

complete a recuperation as has been noted; unless a re-solution of the

erystals and recrystallization were supposed, in which case it may be

argued that such a process would require an abnormal solubility of small

crystals when compared with large. In a normal specimen, re-solution

and recrystallization are undoubtedly going on, strengthening the struc-

ture, and the large crystals are growing at the expense of the small. If

small crystals preponderate at seven days’ age, resulting in a weak mass,

it is necessary to postulate a comparatively high solubility of the small

erystals in order to arrive at a normal strength at 28 days. This, while

by no means impossible, is not probable.

Turning to the specimens kept continuously in cold water, it would

seem that, although the first chilling should show severe effects, as it did,

there should not be such a falling off in the rate of hardening, if the

crystallization be progressive. It is quite possible, however, that crys-

tallization at this temperature is not favored, and that the total number

of binding crystals of calcium hydroxide is therefore less than at normal

temperatures.

QUANTITY OF WATER AT FIRST ADDED

Size of cement particles——Other factors being equal, the amount of

cement rendered inert by the action of water is proportional to the per-

centage of fine particles. This is an absolute condition and presupposes

free access of water to every particle. Needless to say, in practice this

condition is seldom realized, except approximately in laying concrete

under water, or in the careless use of an excess of water in mixing, or in

protracted mixing.

In the use of a very fine cement, then, if the proper proportion of

water is added, the mixing time carefully regulated and proper precau-

tions taken in depositing, the influence of texture upon the strength of

the mass occasioned by the action of water is reduced to a small quantity,

190 ANNALS NEW YORK ACADEMY OF SCIENCES

by virtue of the greater hydraulic activity of the fine particles, increasing

the impermeability, as will be shown, and the confining therefore of the

action of the excess water to a narrow zone. The bulk of the cement will

be properly hydrated in spite of the fineness.

The investigation of the effect of the size of particles due to the action

of water thereon alone is not feasible, because no satisfactory measure of

laitance formation, except the strength of the mass, has been devised.

The measure of the strength would be unsatisfactory, since the propor-

tion of fine particles affects the strength in other ways than through the

formation of laitance, as has been pointed out in a previous communica-

tion.

From a study of the hydraulic properties of reground cement, Spack-

mann and Lesley conclude (93) that only the very fine flour in cement,

that portion not measured by the present tests using sieves, reacts when

gaged with water and gives strength. It is difficult, of course, to draw a

sharp dividing line between active and inactive material in cement, al-

though it must be admitted that the greater part of the coarse material,

even though it be of the same chemical composition as the fine, has little

or no cementing value and serves mainly as a filler.

Suitable fractional separation of the portion of cement passing the 200

sieve, by air-elutriation or other method, should with careful study be a

valuable guide to the most efficient mechanical composition. Experi-

ments upon the first method of separation are recorded by Peterson (71),

and a scientific method of fractional elutriation using an inactive liquid

has been worked out by Thompson (100). Much should be gained by

the application and development of these methods. The influence of the

size of particles of inert material added to the cement is also of great

consequence, and a proper mechanical grading of the sand used in mor-

tars is recognized as vital. The presence of clay in this sand, or the

addition of clay alone to cement, come under this category, and have

occasioned a great deal of discussion (8, 32, 33, 110).

A comparison was made of the permeability of 1:4 mortar of Portland

cement, when used in its ordinary condition, and when screened through

a number 200 sieve.

PACINI, METAMORPHISM OF PORTLAND CEMENT 191

TABLE 3

Permeability of 2-inch Cubes, Age 28 Days, Subjected to 80 lbs. Pressure

Grams of water passing

‘Temperature per hour Nuniboase

Cement of water. — testa

Deg. Fahrenheit

Unscreened Sereened

EN Se Cee See eae RE 66 68 22 By) 5, 6

(SRA cere aoe 68 68 25 2 6, 6

Oc tte, ae eae 68 68 29 Trace 6, 6

Dre 68 68 331 81 5, 6

15r , Serge ey eae eee ee 64 64 rl Trace 5, 6

Ry las: 64 64 31 2 6, 6

CG ja ae 64 64 5 0 3, 6

Heys as. ote SES 68 72 6 Trace 5, 6

Me ye, 68 68 71 0 6, 6

ANY CLE s Goan OM TEC Oa een omer 61 13

The marked decrease in permeability resulting from the use of finer

cement in mortar demonstrates that in impermeability, as in strength,

the finest particles are the most active factors.

Mechanical agitation when water 1s added.—Increased working should

weaken a cement after a certain maximum point is passed. In order to

establish this point, the effect of prolonged working was investigated.

It was necessary to use a mix of fluid consistency, in which, for obvious

reasons, the final set would not under normal conditions take place dur-

ing the time over which the experiments were extended.

Two grouts were employed: one in which cement was mixed with 50

per cent of its weight of water, and one in which an equal weight of

water was used. The different tests were run respectively for periods

from one minute to five hours, and they were mixed in a motor-driven

stirring machine of the type common in chemical laboratories.

After the stated period of stirring, the grouts were poured into glass

tubes and kept in a damp closet for the twenty-eight-day period. Cylin-

ders exactly two diameters high were cut from the specimens and crushed

in the.compressing machine, two cylinders being crushed for each period,

and the average of the compressive strengths being recorded.

192 ANNALS NEW YORK ACADEMY OF SCIENCES

TABLE 4

Twenty-eight-day Tests of Grouts Mixed for Varying Lengths of Time

Compressive strength, pounds

per square inch, average

Duration of mixing ss See ce EUs

50 perct grout /100 per ct.grout

URSIMITNONC vest et or tess ses. cides ence ease 5240 3095

ND TNUTNUTTE Sh. ss o-< ccvsypato se eas eos 9045 4725

POMMMTUMULESe sels se nea gee See: 9710 4955

APO OUI eestis See ease ata mele tonne eerie 5879 4840

POIVOUNS a narcon Eeoiane ae ree ee eee 6075 4320

ENO UNE Sreatenciecveteec aisaeiceee Rare eae 4775 4538

The effect of mechanical agitation, when thus prolonged, is equivalent

to that of the use of excess water—the strength of the cement is pro-

gressively diminished as the working proceeds. It is noteworthy that

the effect is only reached after a certain optimum period is passed. Be-

fore this time, increased working increases the strength. We may con-

clude that there occurs within this period a process which neutralizes the

effect of hydrolysis; and this process is probably the formation of the

network which constitutes the setting.

As will be seen later, the effect of excess water is to reduce the ulti-

mate strength. ‘The effect, then, of mechanical agitation must be to

bring more cement into contact with water and, therefore, to increase

hydrolysis. This is probably accomplished by stripping off the protective

film of gelatinous material which envelops each cement particle when it

comes into contact with water, which film regulates the hydration of

cement and causes it to proceed in a regular manner. This film being

stripped off, the cement is subject to the destructive action of hydrolysis.

Where more water is originally present, the destructive action 1s sooner

attained, as will be seen by comparing the 100 per cent grout with the

50 per cent. Evidently, the setting process proceeds best at high concen-

trations, when the amount of water is low. This may be so regulated

that the setting process will not take place at all, by using a large excess

of water and much mechanical agitation, as has been repeatedly observed.

by the writer. :

Setting time of cement in laboratory ar and im damp closet.—The

standard specifications for setting-time tests call for storing the specimen

in the damp closet, whereas the tests as generally conducted in most

laboratories are made in the open laboratory air. A series of experi-

ments was made, for the purpose of noting the deviation from standard

results caused by this departure from the rule.

PACINI, METAMORPHISM OF PORTLAND CEMENT 193

TABLE 5

Setting Time in Laboratory Air and in Damp Closet

Time of set in minutes

Cement Laboratory air Damp closet

Initial Final Initial Final

RS ie oak See eet ang 255 37d 300 435

VEILS, See oid ates eas ane ae 120 360 300 450

EOE. ieee a as ae ca 300 420 360 480

“Soe atie eee eee 240 360 285 420

NA ssmita rene dco ntct ; 240 390 250 450

From these results, it will be seen that setting in a relatively dry

atmosphere takes place in a shorter time than in a damp one; also that

the setting time is more uniform under conditions of high atmospheric

humidity.

At the same temperature, evaporation takes place more rapidly in the

former case; and allowing a cement mix to stand in such a position that

evaporation of the mixing water may readily take place is practically

equivalent to the use of an insufficient amount of mixing water.

Effect of excess of mixing water on strength of concrete.—Concrete is

often mixed so wet that, as it is filled into forms to a depth of several

feet, the water rises above the concrete and throws out considerable lai-

tance from the cement. The ease of mixing and placing very wet con-

erete is the constant incentive for its use. This practice, however, is

followed by a great deal of deterioration of the concrete in strength.

The strength rapidly decreases with the increase in the quantity of

_ water used in mixing. The visible effect of this weakening is the forma-

tion of laitance, which has little or no setting power or strength, and

which represents the loss of an active part of the cement, since, as is

recognized, the finer parts are more hydraulically active.

Tests were made by mixing concrete at normal consistency and shovel-

ing one-half the batch into a tank containing three to four inches of

water, the depth of concrete being about four inches. The water rose to

about an equal depth above the concrete. In test No. 1, the concrete was

allowed to settle in water four inches in depth for 30 minutes, when the

excess of water was siphoned off and the remaining material poured into

molds. In test No. 2, the depth of water in the tank was three inches,

and the water was siphoned off immediately while in agitation. In test

No. 3, the same process was repeated, except that the depth of the water

194 ANNALS NEW YORK ACADEMY OF SCIENCES

was four inches, and the concrete used was somewhat leaner. Test No. 4

represents the direct qualitative effect of the addition of an excess quan-

tity of mixing water without subsequent handling. All specimens were

cylinders six inches in diameter and 12 inches high. The remainder of

the batch of concrete in each case was poured directly into molds, and the

specimens were broken at 28 days. The amount of cement lost was

roughly ascertained where possible by filtering the siphoned water and

weighing the amount retained on the filter.

TABLE 6

Effect of Excess of Mixing Water on Strength of Concrete

Strength at 28

No. of . Per cent

Test No. = Proportions days, pounds

specimens of water per sq. in.

Per cent of

cement lost

1240

760

8.2

0.3

»Specimens shoveled into water as described.

Evidently, then, the mere presence of an excess of water is sufficient to

produce the weakening effect, independently of any actual removal of

- cement from the concrete. As may be seen from Nos. 1 to 3, the leaner

mixes suffer the greater deterioration in strength.

Effect of eacess of mixing water on permeability of concrete-—A par-

allel series of tests upon the permeability of concrete treated with an

excess of water was made, in which the correspondingly numbered speci-

mens were treated in the same manner. The cylinders cast from these

batches were eight inches in diameter and six inches in length, and were

cased in the standard manner for permeability tests. Three specimens

were made for each test, and at the age of 28 days were submitted first to

40 pounds pressure for one hour, then to 80 pounds for one hour, without

interruption. The flow recorded is in grams passing during the last ten

minutes of test.

PAOCINI, METAMORPHISM OF PORTLAND CEMENT 195

TABLE 7

Effect of Excess of Mixing Water on Permeability of Concrete

Gaps passing in last

mn en minutes

* Test No. Proportions REE cont of A aareoiine

water

40 pounds 80 pounds

Wend 24 8.2 67° F. 0 0

1 he 74 24 Sie 2ea ania e lercsperceors 479 456

B 2 24 8.2 58° F. 0 0

Be IZ 24 Sere Minne er alae 212 588

3 1: 2.33 25 8.2 56° F. 0 alt

ole 1:2.33 :5. 8.2 60° F. 1814 Not tested

4 1 a jam aaa 8.2 67° F. 38 18

4 ieee 24 MOSSE aes [Utes temas carer 26 80

8 Specimens shoveled into water as described above.

In the foregoing experiments, the decrease in strength and water-

tightness may be referred to the deteriorating influence of excess water

upon the cement (16). It may of course be argued that the more

marked effects obtained in series 1, 2 and 3 than in series 4 are due to

the method of making the tests; that is, that a considerable proportion

of the active cement was actually removed from the body of the concrete

by siphoning off the supernatant water with its laitance.

Effect of excess of mixing water on the strength of neat cement.—

With the idea in mind that the weakening effect was independent of the

removal of cement (1), a further series of tests was instituted, using a

neat cement of good quality. The cement was poured into a series of

glass tubes in which increasing proportions of water had been put, the

tests representing a series of grouts mixed respectively with 50, 75, 100,

150, 200 and 500 per cent by weight of cement of water. The tubes were

shaken for one hour and then allowed to stand for 28 days. The cement

settled into the bottom of the tubes in the order of its coarseness, the fine

nebulous laitance settling last as a cheesy white layer of increasing thick-

ness, as the percentage of water was higher. This layer was carefully

trimmed off in preparing the test specimens.

On breaking out the cylinders from the tubes at the end of the test

period, it was decided to cut each cylinder into two, each exactly one

diameter high, carefully noting the respective position of each in the

tube. On submitting these to compression it was seen that the direction

of difference between the upper and lower layers was not constant, nor

196 ANNALS NEW YORK ACADEMY OF SCIENCES —

was the difference a significant one, so that it was considered wo

to average the strengths.

It will be seen by the table below that, even without actual removal of

any cement, the formation of laitance has a weakening action ge

cement.

TABLE 8

Compressive Strength of Grouts Mixed with Varying Proportions of Water

Crushing strength,

Per cent of water eee pees ineh,

Age, 28 days

50 6855

79 5900

100 4500

150 3430

200 2960

500 1810

The effect of excess of mixing water is therefore seen to result in

decrease of strength as the water increases. Whether the effect is a

permanent one was the next question that presented itself. 'To settle

this point, a new series was undertaken, in which a larger number of

differing percentages was introduced, and in which the resulting strength

at two periods was determined.

The cement was mixed with the stated percentage of water, and

worked for two minutes, the drier mixes upon the table in the usual

fashion, and the wetter mixes merely poured into the tubes and shaken.

Paper mailing tubes were used, 2 inches by 48 inches, treated with

molten paraffin and sealed with paraffined corks, so as to be absolutely

tight. To obviate the effect of possible leakage, the whole series was

stored in damp sand.

Cylinders two diameters high were cut from the specimens at the

stated periods, each cylinder being cut as nearly as possible the same dis-

tance from the bottom, and care was taken to avoid including any of the

soft cheesy top portion, the settled laitance.

PACINI, METAMORPHISM OF PORTLAND CEMENT — 197

TABLE 9

Compressive Strength of Grouts Mixed with Varying Proportions of Water,

Over Extended Period

(Hach result is the average strength of three specimens. )

Compressive strength, pounds per : ;

Percentage of Sy aera Per cent gain in

water strength over

28 days

28 days 3 months

22 7076 7504 6

25 6174 9402 —13

30 4563 6030 32

38 3992 5059 27

50 2991 9312 77

79 2113 4078 93

100 1609 3044 120

150 1270 2379 87

200 1306 2579 97

500 399 1141 186

It is apparent from these figures that the effect of hydrolysis upon the

strength of cement is a reversible one, at least to a certain extent, since

the specimens in which an excess of water was used in mixing showed a

greater recuperative ability at the longer period than the cement in

which the normal amount of mixing water, in this case 22 per cent, was

used.

Upon inspection, it was observed that the three months’ specimens

showed in each case much less laitance than the similar 28 days’ speci-

mens had shown, and it was considered probable that the laitance, in

standing, had adsorbed free lime from the remainder of the cement,

through the activity of the water permeating the mass, and thus reverted

to the original condition of the cement, or an approach thereto. An

analysis was accordingly made of laitance scraped off from the top of one

of the 500 per cent water specimens and thoroughly washed by decanta-

tion. It probably represents a maximum condition in the hydrolysis of

cement.

TABLE 10

Analysis of Laitance from 500 per cent Specimen

As obtained from | Treated with lime

specimen water

SiO ea See hon suet oat 15.28 15.91 ©

CRO Brae sere decease wvevaie a els, okt oneivove 2.28 2.42

PACE) raps cea, estat pcre ccdaiesie si 3.98 9. 82

(OPO) oR ahi yn al rt 26.96 - 86.67

LIVE Oe i cae ct crate ala este oy oie 2.86 1.28

SOM rics civscen vavsmicrs ie aikee chat 6.47 Ween

COM TCO; Chile even sch whe os 42.17 35.18

198 ANNALS NEW YORK ACADEMY OF SCIENCES

The normal ratio of silica to lime in unset cement may be considered

1 to 2.82. In this material we find the ratio 1 to 1.76. This indicates a

great loss of lime; and it was thought possible, that, by adsorption of

lime, this laitance might regain at least a part of its hydraulic proper-

ties. Accordingly it was digested for several days with lime water at

laboratory temperature, filtered off, carefully washed with distilled water

and dried, as was the previous sample, at 100° C. An analysis showed

the results tabulated in the second column. The ratio of Si0, to CaO

had changed to 1: 2.30.

Besides direct metathetical reactions between the components of ce-

ment and the water solution which always surrounds a mass of hardening

cement, adsorption of various materials from this solution is unques-

tionably always going on. Were the fine particles of cement inert chem-

ically, this would still take place, by virtue of the enormous total surface

which they must present. Clay, it has been demonstrated, has the prop-

erty of adsorbing ions of CO, from solutions of carbonates, and of Cl

from solutions of chlorides (10).

The laitance then may, by adsorption of calcium hydroxide given off

from the cement adjacent to it, recover some of the lime lost by it.

Whether the lime adsorbed restores the original status of constitution is

of course mere speculation. The trend of the strength tests shows that

this is probably not so, but that the adsorption is not entirely a reversion

of the hydrolytic reaction; in other words, that “drowned” cement will

probably never recover and attain to the strength it would have had with

proper hydration. |

Effect of the presence of clay and dissolved substances.—It is apparent

that if the decreased strength be directly referable to the action of the

excess water upon the cement, any means of preventing the access of

excess water should prevent, if only to a degree, the destructive action.

The colloidal nature of clay (6) has been utilized in the water-proofing

of concrete, the principle of its action being the formation of continuous

gelatinous films throughout the structure, which prevent the passage of

water. Although the same problem is not presented in a grout that exists

in finished concrete, it is probable that some blanketing action might

occur upon the addition of clay to the mixed mass.

The point was investigated. To correct for the effect of absorption of

part of the mixing water by the admixed clay, a consistency test was

made upon a sample of cement to which 10 per cent of clay had been

added, and it was found to require 4 per cent more water than the same

cement used neat. :

The clay mixes were accordingly gaged with 4 per cent more water

PACINI, METAMORPHISM OF PORTLAND CEMENT 199

than the corresponding neat cement mixes, and the following series of

compressive tests was made:

TABLE 11

Effect of Clay upon Destructive Action of ‘Eacess of Mixing Water

(Average of two tests at 28 days)

Neat cement Sere ace

Compressive Compressive

Water, ; Water,

Beef ange ence ECE a gpereent (| ete ete poulndl

50 5782 54 1282

75 3134 79 1328

100 2273 104 2577

150 1896 154 2156

200 1381 204 1320

900 514 504 No strength

developed

If the action of saline solutions upon cement is to accelerate the hy-

drolysis of the latter, it would appear that the destructive action of excess.

water would be accelerated by the presence:therein of saline substances.

in solution; also, it is legitimate to expect that the addition of clay

restraining the hydrolysis due to excess water will in this case exert a

similar influence.

The following experiments, parallel to the foregoing ones, elaborate

this point :

TABLE 12

Effect of Clay upon accelerated destructive Action of Mixing Water Containing

5 per cent of Magnesium Sulphate

(Average of two tests at 28 days)

Cement, 10 per cent of which was

Neaticemenp replaced by a fat clay (dried)

5 per cent solution Compressive 5 percent solution Compressive

of magnesium strength, pounds of magnesium strength, pounds

sulphate, per square sulphate, per square

per cent inch per cent inch

50 2196 54 2774

75 548 79 1608

100 1512 104 No strength

150 556 154 a is

200 ' No strength 204 re es

500 No strength 504 os ee

200 ANNALS NEW YORK ACADEMY OF SCIENCES

From these two series of experiments, it is qualitatively apparent that

the presence of clay does prevent a certain amount of hydrolysis. From

the first series, it is seen that this effect only begins to show itself as

higher percentages of water are present, which would indicate that the

clay may have taken up much more water than the constituency test

revealed, and that, in the relatively drier mixes with clay, the cement

suffered in strength because of insufficient water. On the other hand,

experiments at this laboratory in which clay was used, replacing up to

10 per cent of cement in normally gaged material, showed that no signifi-

cant decrease in strength was thereby obtained ; hence the loss in strength

in the 54 and 79 per cent grouts cannot be due to this cause.

_It is more probable that the colloidal nature of the added clay is

brought into play more effectively at the concentrations in which in-

creased strength is observed, and that the latter is due to the coagulation

of the clay by electrolytes adsorbed at this optimum concentration.

The same result would obtain where additional saline material has

been added to the mixing water, as in the series where a 5 per cent solu-

tion of magnesium sulphate was used. The clay here prevents the accel-

eration of hydrolysis by the magnesium sulphate through adsorption of

part thereof, and possibly by coagulating, forming an impenetrable bar-

rier to the further action of water upon the remainder of the cement.

QUANTITY OF WATER THAT MAY SUBSEQUENTLY COME INTO CONTACT

WITH THE SYSTEM

Permeability—The solvent effect of water coming into contact with

cement structures is best studied by the permeability test. This consists

in forcing water through a mortar or concrete at a known pressure and

observing the amount of leakage through the specimen. In detail, the

specimen is generally made up in the form of a cylinder, and this is

cased with a thick coating of neat cement on all sides but the bottom.

The water, under pressure, is applied on the full cross-section of the

specimen and forced through, dripping from the bottom, whence it may

be collected. ;

With neat cement, of course, this method is inapplicable, because of

the density of the material and the consequently enormous pressure nec-

essary to force water through it, and moreover because of the mechanical

difficulty in confining the water strictly to a passage through the speci-

men. ‘The specimens tested, then, are lean mortars and concretes.

Although this test is designed to ascertain the resistance which these

materials offer to the flow of water, it is evident that this resistance is

not a constant quantity in the case under consideration.

PACINI, METAMORPHISM OF PORTLAND CEMENT 201

The temperature and pressure of the percolating water being constant,

the flow is diminished by cementing and clogging, and increased by ero-

sion and solution; the quantity of water flowing through the mortar or

concrete therefore is a function of the balancing of these processes. .

Cementing may result from deposition of material originally in solu-

tion in the percolating water, or dissolved from one portion of the

structure and deposited in another.

Clogging, similarly, results from material originally in suspension in

the percolating water, and deposited in the pores of the concrete, or from

material eroded from one part of the mass, either mechanically or as a

result of solution of the attacking portions, and deposited in another

part.

Erosion per se is a negligible factor; that is, the flow of pure water,

carrying no suspended matter, will have very small mechanical effect

upon an insoluble material. When the water is armed with suspended

matter, however, its corrasive effects become proportionally magnified.

Solution is the most important factor in the process of percolation.

Following the order laid down by Van Hise for natural rocks (104, p.

536.), the basic materials removed are, firstly, the alkalies and, secondly,

the alkaline earths, in the order calcium, magnesium. Since the alkalies

exist In cement in the proportion of a little over one per cent and are

not essential to the hydraulic properties or the strength, their solution

is a matter of little consequence, except in that it may result in the for-

mation of solutions which react upon the lime compounds and render

their solution more easy of accomplishment. This reaction has been

considered elsewhere. The removal of magnesium compounds proceeds

at a lesser rate, although there is a greater percentage of them present ;

and their removal, in the main, may be dismissed as insignificant.

Since more than half the weight of fresh cement consists of lime, and

since the strength of cement depends for the greater part upon calcium

hydroxide, whether crystalline or adsorbed by colloids, the removal of

calcium hydroxide from set cement is the factor of the greatest impor-

tance. Considering its solubility in pure water, the reversion of the

hydroxide to the crystalline form tends to diminish its solubility, or

from the other standpoint, its adsorption by a colloid tends to remove it

from the solvent action of water. Unfortunately, however, it must be

borne in mind that without exception, cement structures are nowhere

subject to the action of pure water alone. From rain water, with its

appreciable burden of dissolved gases and atmospheric salts, to the

water of the ocean and the more heavily laden rock and mine waters,

concrete structures are everywhere in contact with saline solutions of

varying concentrations.

202 _ ANNALS NEW YORK ACADEMY OF SCIENCES

The effect of solution in percolation, then, is to a small degree de-

pendent upon the solubility of the components in pure water. This effect

diminishes as time goes on, because of the reversion of the soluble ma- .

terial to a less soluble form and because of the protection afforded by the

insoluble portions of the system decreasing the exposed area of soluble

material. The washing away of these protecting films will of course

neutralize the second factor. The increased solubility of the components

of set cement in solutions of various electrolytes is the more important

element in percolation. Even a very dilute solution may have tremen-

dous total solvent power, when the time element is considered. In fact,

it may be that the action of a dilute solution will on the whole exceed

that of a concentrated solution, by reason of the greater cementing and

choking action of the latter, tending to diminish the quantity of water

that may come into contact with the soluble portions. A dilute solution,

therefore, with its more insidious attack, is probably more to be feared in

the end than the strong brine.

Observation of the behavior of concretes and mortars during the per-

meability tests gives a clue to the balancing of these processes, whether

there is a preponderance of cementing and clogging on the one hand, or

of solution and erosion on the other. Attempts were made, in the experi-

ments noted below, to study chemically the reactions involved, by peri-

odical analyses of the percolating water. To this end nearly four hun-

dred complete analyses of the effluent water were made. Upon tabulation

of these it was observed that any deductions based upon them would be

inconclusive, as the chemical composition of the effluent water repre-

sented one of a great number of variable factors that might occur at any

point either within or without the concrete. The single qualitative

generalization, that lime was removed from the cement at a diminishing

rate, is the only permissible conclusion from the analytical data.

The original purpose of these tests was to ascertain the suitability of

various aggregates for use in concrete, with reference to their stability

in the presence of percolating water. At the conclusion of the series, it

was found that the effect of water upon the various aggregates was prac-

tically negligible, during the period of observation, and that the action

had been confined to the cement of the mortar. The aggregates had been

protected from the action of water by the cement, it being probable, how-

ever, that a continuation of the tests would have revealed the action of

water upon these rocks, when the protective influence was removed.

A series of sixteen aggregates was used, in as many concrete specimens.

Since it is not the purpose of this report to discuss the relative suita-

bility of these materials for concrete construction, but only to consider

PACINI, METAMORPHISM OF PORTLAND CEMENT 203

the action of the water upon the cement, two cases alone will be con-

sidered.

The rock was crushed and screened for each experiment to the same

average effective size, corresponding to the following mechanical analysis:

TABLE 13

Mechanical Analysis of Aggregate used in Permeability Tests

Sieve ee ve Per cent passing

1% 1.89 100

1% 1.58 94

1 1.02 a9

x .78 32

24, 59 21

7 .48 16

3 .30 6

4 .22 0

The sieve ratings are based on diameters of spheres of equivalent vol-

ume to the largest sized stone particles that will pass.

The fine aggregate was crushed quartz, the standard sand formerly

used for cement testing, passing the No. 20 and retained on the No. 30

sieve. The cement used was a standard Portland of high quality.

The specimens were made in the laboratory’s standard form for per-

meability test, cylinders eight inches in diameter and six inches in length,

the proportions used being 1:3.5:6, this being found the richest mix

practicable to secure the porosity required for the test. They were cased

in neat cement, and connected suitably for subjection to the pressure of

the city’s water mains. Each specimen was protected from the direct

flow of the water by a layer of one inch of clean coarse sand. The average

pressure for the period of observation (52 weeks) was 22 pounds. The

determinations of the rate of leakage were made weekly at first, and later

every two weeks until the end of the test.

The data appended below represent observations on the rate of percola-

tion of water through two of the specimens which present the greatest

interest from the standpoint of this paper, this flow being recorded in

grams passing in ten minutes. The aggregate used in one specimen was

a hardened neat cement, crushed to the size stated, and used in place of

the rock generally employed in concrete. The parallel specimen selected

for comparison was one in which the aggregate was a crushed granite,

which showed a low solubility in hydrochloric acid (2.66 per cent dis-

solved in one hour’s treatment with 1:1 HCl).

204

Temperature records of the percolating water were not kept, since these

tests represent a part of a larger series in which this would have been

impracticable. The other aggregates tested showed results from which it

was quite difficult to draw any legitimate conclusions as to the relative

suitability of different rocks in concrete subjected to these conditions.

Concretes containing different aggregates.—A series of tests on con-

cretes made up of different aggregates but with the same cements gave

results which may be tabulated as follows:

Flow in Grams of Water passing in 10 Minutes through Concrete Specimens

TABLE 14

ANNALS NEW YORK ACADEMY OF SCIENCES

subjected to continuous Water Pressure for 52 Weeks

Pressure,

Time pounds per

square inch

24 hours 25

1 week 2

2 weeks 22

Ope 20

AT? 25

Satie 22

Ce =a 20

"/ ce 20)

Sibi 25

LO: ae 25

Date 26

MA Sabte 24

Gy 24

alfeyen ee 20

20 17

2 aS 20

Pye ee 17

26ic0e- 22

DBR es 26

3007 = 22

Bede alr 20

Sy ie 19

ai 26

Sos 26

40 *“ 23

AO oe 25

44 * 22

Age 21

AG WIE 25

50 20

SDA Race 20

Month

January....

February ..

March.....

August.....

September.

October....

November..

December. .

January....

Grams passing in 10 minutes

Conereve with Concrete with

aggregate of

erushed aggregate

of crushed

oat comonte| = Eee

2111 60

836 31

662 26

626 40

570 60

603 62

530 50

1295 38

1127 40

1310 45

870 25

997 36

985 28

973 32

639 20

792 40

731 36

802 43

800 49

781 46

763 50 ©

115 Trace

105 2s

107 2}

110 3

93 oD,

75 9)

70 3

80 10

73 ee:

78 if

PACINI, METAMORPHISM OF PORTLAND CEMENT 905

Comparison of these two sets of figures indicates that the cement of the

concrete is more attacked than the aggregate. In fact, the flow obtained

in this specimen was the highest but one of a series of sixteen, and the

total lime content of the effluent water was also the highest but one.

The visible effect upon examination of the interior of the specimens

was a bleaching of the mortar, with evident solution of the cement. The

original percentage of lime in the mortars was 12.8. Analysis of mortar

from the granite specimen showed a content of 4.8 per cent, indicating

that nearly two-thirds of the lime had been dissolved out. Further evi-

dence of the loss of lime was found in the heavy white crust which

formed on the exposed bottoms of the concrete specimens during the test.

Small stalactites, quite soft to the touch, were abundant. The quantity

of this deposit was not visibly different in the different tests.

The calculated loss in lime of the mortar was greater than the loss

computed from periodical chemical analyses of the effluent water, and

this is due to the fact that much of the dissolved lime was deposited upon

the bottoms of the specimens as the stalactitic growth above mentioned.

There was no evidence that suspended impurities in the water had

been carried into the interior of the concrete, and it is therefore supposed

that the one-inch layer of sand by which the latter was screened from

the direct flow of the water was an efficient filter for the purpose. The

clogging action resulting from this source may therefore be dismissed as

negligible.

It may be concluded from these tests that concrete of this density tends

to protect itself automatically from the action of percolating water, so

that, for the period investigated at least, the flow tends to diminish to a

minimum. The action of the water seems to be confined to the cement

of the mortar, leaving the aggregate relatively unaffected.

It is evident that, notwithstanding the utmost precaution in mixing

concrete test specimens, wide differences in permeability may obtain in

specimens mixed under the same conditions of handling and by the same

workman, owing to structural differences in the resulting mass. How-

ever, the results obtained are fairly comparable.

_The most sensitive test for the internal changes which the concrete has

undergone during percolation is the resulting strength of the concrete.

Concretes containing different cements——A series of tests was under-

taken in which the specimens were made up in the same proportions,

1: 2.5: 6, using in each specimen the same coarse aggregate, a crushed

eranite, and the same fine aggregate, a standard quartz, but using differ-

ent brands of cement. The specimens were stored in damp sand for a

period of 28 days, then subjected to continuous water pressure of about

206 ANNALS NEW YORK ACADEMY OF SCIENCES

25 pounds for a period of 11 months.

the specimens :

: Percolation through Concrete Specimens

Parallel specimens were stored in

damp sand during this period and allowed to attain their full normal

strength. The table following shows the leakage and final strength of

TABLE 15

Months of percolation

Brands of cement and grams of water passing in 10 minutes

Compressive strength of specimens

at the end of period

Compressive strength of untreated

specimens, pounds per square inch..

Loss of strength through percolation

A B C D 1D) F

146 286 o 164 76 230

155 125 22 179 16 82

a6 70 90 167 11 85

37 47 52 161 11 82

U2 28 37 65 a 45

71 12 31 15 17 39

68 28 3 11 26 33

a7 46 14 6 16 21

40 43 atte 2 5) 11

18 oie 10 “5 a 14

8 13 10 ‘1 2 19

TABLE 16

Comparison of Strength before and after Permeability Test

A B C D EK F

avs ae cave eae ane 770"| 490 640 890 750 | 8590

1080 | 1210 | 1230 | 1125 1220 1090

29 60 A8 21 39 46

7 One specimen crushed.

(DETZCemt hs eee are ee ener

Other results are average of two specimens.

Effect of the direction of flow through concrete.—Concrete seems to

offer less resistance to the flow of water when the direction of the flow is

parallel to the bed than when at right angles to it. A test covering this

point was made with 8-inch cubes of concrete of the proportions 1: 4: 14,

fine and coarse aggregate being a standard crushed bluestone.

PACINI, METAMORPHISM OF PORTLAND CEMENT 207

TABLE 17

Rate of Flow in Gallons per Square Foot per Hour under 20-inch Head

Age of specimens, 67 days. In specimen parallel to In specimen perpendicular

Temperature of water, 64° F. bed to bed

NSGe2emMIMUteS 2 sce se. oe 740.96 164.14

2d oes Ra ote ess ede ca" 585.28 159.54

3d TBE Teen A USI, 2 alee, 636.31 163.49

4th OM esr syt et Mer ca 539.53 158.3 |

5th USI Mea aa A Nt a 549.10 157.93 |

Specimens immersed 24 hours, then retested :

“US A ONO) Se Seee nes ee 665.38 182.46

2d CO AON th ied Ba We Oe Bee 642.77 177.54

3d AO TO tee EE eae Ne a 662.80 177.54

4th Dis SIT eer ame Reena 641.15 177.06

5th us eRe nner hs 659.57 173.67

In denser concretes, this effect was not found so marked. It will be

noted that after storage following the first exposure to the effect of per-

colating water, these specimens appear to offer less resistance to the flow

of water. This may be due to the fact that in lean concretes the propor-

tion of capillary and subcapillary voids is smaller and that of super-

capillary voids greater, and that cementing and clogging actions, which

have their greatest effect in capillary and subcapillary passages, are not

so effective.

The greater flow along the bedding planes has been observed in the

case of rock, and is in all respects a phenomenon of the same nature. In

the case of a stratified sandstone cited by King (51), the reason is ad-

vanced that no more water can pass the more open layers, when advancing

across the bedding planes, than was able to pass those of the closest tex-

ture; whereas when the flow is along the bedding planes, each particular

stratum carries water in proportion to the coarseness of its texture,

uninfluenced by any other.

In the case of water percolating into a concrete tunnel this would

tend to emphasize lateral percolation, and in the case. of disintegration

would exercise, in general, a localizing influence. It is not to be as-

sumed that this is a rigid rule, inasmuch as a large number of factors,

evidently, may neutralize this influence.

From these considerations, it will be seen that the solvent effect of

water upon set cement is of high importance in considering the perma-

nence of concrete structures, and that this solvent effect tends to diminish

as the set cement ages. This is not the only way, of course, that water

208 ' ANNALS NEW YORK ACADEMY OF SCIENCES

may afterwards affect the metamorphism of cement. It has been pointed

out by Goldbeck (43) and by White (108) that the expansion or con-

traction of concrete depends upon whether the concrete remains wet or

dry, and that the strains caused by alternate wetting and drying of con-

crete may be a more fruitful cause of cracks than temperature changes.

The presence of an optimum quantity of water is necessary, however,

so that the proper reactions take place in the mass of setting cement, in

order that the strength may increase normally.

QUALITY OF WATER AT FIRST ADDED

Compressive strengths of neat cements gaged with various solutions.—

A normal Portland cement was mixed with the proper quantity of

water (21 per cent by weight) in which was dissolved, in the different

tests, varying concentrations of the salts indicated in the subjoined table.

he cement was worked for one minute, and the plastic mass was tamped

into glass cylinders approximately one inch in diameter, with the utmost

precaution to avoid all air bubbles and at the same time to subject all

specimens to the same pressure.

TABLE 18

Compressive Strengths of Neat Cement Mixed with Solutions of Various Salts

(Age of specimens, 28 days. Average of two determinations)

Mikes Pounds per | Gain or loss,

square ineh per cent

il. Distilled qyater2 4-5 fo aerate ee eee eter 7330

2. 25% rock water® diluted with distilled water.... 6340 —14

3. 50% do. Ge 6495 —11

4. 75% do. ee, 6870 — 6

On AROck water alone: ...¢ ascertain 5605 —23

6. 2% sodium chioride.solution4.. 4.2. ee. eee ~ 6675 — 9

7. 4% GO. AL © a tgs Sag te Pees las eo ae ea 5815 —21-

8. 6% do. Fae ie Sar Aka eek Ree cite Ue Oma 5065 —31

9. 8% (a (Gea RNa Ca Ket Ree grees Fos eat pacchore es a 4215 —43

10. 10% Oe: Sepa linus Say ar outer eee Ce ee 5285 —29

1]. Saturated solution of calcium sulphate (+ 0.2%). 7025 —4

12. 0 2% solution of calcium chloride............... 6960 —5

13. 0.2% solution of magnesium sulphate............ 6680 — 9

14, 0.2% solution of magnesium chloride.. ane 59995. —23

15. Equal parts of 11 and 12 (CaSO, and CaCl, ae liad 6565 —10

16. Equal parts of 13 and 14 (MgSO, and MeCl, Nera 7355 +0.6

17. Equal parts of 12 and 13 (CaC), and MgSOQ,)..... 5810 —21

18. Equal parts of 11 and 14 (CaSO, and Mg(l,)..... 6200 —15

§ This water contained: CaO, 1177 parts per million.

MgO, 226

SO;, 408

Cl, 4360

PACINI, METAMORPHISM OF PORTLAND CEMENT 209

The glass cylinders containing the cement were then stored ina damp

closet for 28 days, when the cylinders were broken out, and two speci-

mens, each exactly one diameter high, cut from each cylinder. These

were put into water for a few hours, so that they might be in the moist.

state when crushed. The cylinders were kept in the damp closet instead

of being stored under water, to avoid leaching out the salts contained in

the mixing water, thus obtaining the maximum effect of the dissolved

salts.

It will be noted that there is a decided loss of strength in all but one

ease (number 16). This particular case may be explained by the prob-

able formation of an oxychloride, by the magnesium chloride and the

magnesium hydroxide liberated by the action of the magnesium sulphate

upon the calcium hydroxide of the cement. The oxychloride formed

from these two materials has a tensile strength far superior to that of

Portland cement itself, and its presence probably counteracted the de-

structive action of the salts upon the cement. It is probable, however,

that, at longer periods, this increase would disappear and become a de-

crease. Otherwise, the presence of saline matter dissolved in the mixing

water seems to have a decided deleterious effect upon the strength of

cement. ‘This point is of marked importance in construction, inasmuch

as the problem of mixing water is often solved by using the water nearest

at hand, without inquiry into its qualities.

Tt is the custom to specify that the water used in mixing concrete shall

be free from oil, acid, strong alkalies or vegetable matter (77) ; but such

_ a Specification does not cover the case in point, and the presence of large

quantities of dissolved salts in water used for construction is easily over-

looked. In concrete construction, it is of the utmost importance that the

water which may be used in mixing be additionally subjected to such

tests as will reveal either its mineral content or its action when mixed

with cement and possible subsequent attack thereon.

The action of sodium chloride appears to be nearly directly propor-

tional to the amount employed. This salt is used in mixing water for

construction carried on in cold weather, in order to prevent freezing of

the deposited concrete. Its effect upon the strength of cement, if used in

excessive quantities, is, as has been shown above, likely to become a seri-

ous matter. Under the conditions of construction which generally pre-

vail, however, much of the salt may be leached out of the mass. The

results above represent a condition of maximum attack.

Dieckmann (25) recommends the use of from 1 to 2.5 per cent of salt

for concrete to be laid in cold weather, but states that percentages larger

than this cause a marked decrease in the strength.

ji) ANNALS NEW YORK ACADEMY OF SCIENCES.

Effect of gaging with various solutions upon the strength of mortars

afterward stored in water.—The above tests do not show, of course, a

normal condition, since no water came into contact with the cement

after it had set. Working with more porous material, a 1:3 mortar, so

that in storage a heightened subsequent water action might take place,

the following results were obtained :

TABLE 19

Effect of various Salts dissolved in the Mixing Water, wpon the Strength of

1:3 Mortar

(Sand, screened Cow Bay. Specimens stored in damp closet for 24 hours, then

continuously in water for the rest of period)

Compressive strength,

pounds per square inch

Number of

specimens

7 days 28 days 3 months

815 1475 2600 1

1005 1185 1805 3

945 1310 2170 3

1010 1520 2420 3

885 1240 2100 i

910 1625 2145 2,

ww we ewe wv

CD OD HE Od HI OD GD CO CO CD GO KD OO

G9 9 OO bo bD OD 09 CO CD CO EI CD

865 1410 2115

935 1595 2710

2% (6 Co RCNA he. Va eM TEM os 1090 1580 2500 sie

i ssolution Of BESO pa. s-eeseRe 930 1605 2670 135

2% OE 1 fo S20, Gee eee aioe 840 1480 2710 2.9.

lZasolatonotNa@late.ce ere ener 1105 1385 2000 3,9,

2% GOs oA ee EE eee eae 1000 1035 1685 2,05

Tensile strength, pounds per

square inch

7 days 28 days. | 3 months

NUCH Ta" ores tie part pk Le ee ara ie Sin 179 272 326 5,6,6

1% solution of Al, (SO,),.......... 208 272 321 6,6,6

2% HOSS) 5S OSB Fa bea ete eae 193 262 340 4,6,5

1% solution of Na,SO,........ Gia 216 290 304 6,6,6

2% GOR oot adie are ee ene ears 205 300 343 4,6,6

1% solution of MgSO, ........2. 02 6: 194 260 317 3,6,6

2% CG LORIN Seca © pei gtaninn Tune 185 246 283 46,6

1% solution of ZnSO,.............. 126 263 315 2,0,6

2% QE! fact eee goer 205 272 319 3,6,6

1% solution of FeSO,.............- 201 266 314 5 ,6,6

2% (0 (On Celene Reena tc ARS 184 258 311 5,6,6

1% solution of NaCl............... 211 261 310 6,6,6

2% Os pra soe crease pee Weck dae 224 209 310 5,6,5

PACINI, METAMORPHISM OF PORTLAND CEMENT 911

The general conclusion that may be drawn from these values is that

the effect of electrolytes in the mixing water, when the cement is after-

wards subject to immersion in water, is to increase the strength at the

early periods (7 and 28 days), but later to depress it (15). In general,

the more concentrated solutions give a greater depression of strength.

The early increase in strength is probably due, in the presence of an

optimum quantity of water, to additional cementing or void-filling ma-

terial precipitated in the pores of the mortar by reaction between the

added electrolytes and the solutions resulting from the action of water

upon cement. This deposited material may, in its later history, revert

to a soluble form and be washed away, leaving abnormal voids, or else

. in its growth may disrupt the cells it occupies, in either case reducing

the strength.

Effect of gaging grout with rock waters.—In grouting deep tunnels,

the question has arisen as to the advisability of using the rock water at

hand when fresh water was inaccessible. The water available in the

instance in hand was an effluent from a shale bearing a small proportion

of pyrites, and when it issued from the rock face it contained a quantity

of dissolved hydrogen sulphide. As none of the water was immediately

available for a laboratory test, an artificial mixture was made up, in

which the quantities of dissolved salts and hydrogen sulphide occurring

in the natural water was purposely exaggerated, to obtain accelerated

effects.

TABLE 20

Analysis of the Artificially Mineralized. Water

Parts per million

IETS iptrstevrate vatettostsna etait soderter eels titotes toys Rosh rateita tal Me ltite 891

AOI Gi ee trate ttta ater RR ORE I 1764

IM EXO) Sala pick o Oa OiGs 1050S COSMOS CRS areca sc 1461

S Olpreyseot oa cronnnesr dare atie dias cvcrinseisie ts auselaa 1948

(Ol aes saa encieites Sr itire ens ogee pe eRe eae aaa Ard 2920

A grout was made up according to specifications, using a normal Port-

land cement, and Cow Bay sand with 100 per cent passing 10 sieve, 75

per cent passing 40 sieve; in the proportions 1: 1144 with 35 per cent of

liquid. The wet mix was poured into glass cylinders, kept 24 hours in

air until set had developed and immersed in water.

Four sets of three specimens each were made, the first set mixed with

35 per cent of distilled water; the second, 35 per cent of the water above

mentioned ; the third, 35 per cent of a 10 per cent dilution of this water,

and the fourth, 35 per cent of a 1 per cent dilution.

No discrepancy was observed in the setting time, as all the specimens

912 ANNALS NEW YORK ACADEMY OF SCIENCES

developed a fair set within 24 hours. The grouts mixed with the undi-

luted sulphide water turned a dark green, but otherwise no change was

noticed in these or any other specimens. Three cylinders one diameter

high were cut from each set of specimens, and, after storing 28 days in

distilled water, were crushed.

TABLE 21

Compressive Strength of Grout Mixed with Different Proportions of Water

Containing Hydrogen Sulphide

(Average of three specimens, age 28 days)

Pounds per

Mixes square inch

DD USEMN OG: WiabOr 55.005 Soe wed vo, cen Sas Gr wale airatotlinte eilote cneiles She tatielte een ne 1424

Undiluted sulphide wateric)j.4.c. acc 0s Soe oes Se ae 1608

10 per cent of sulphide water, 99 per cent of distilled water.... 208

1 per cent of sulphide water, 99 per cent of distilled water...... 1110

Apparently, considering the average of the last three values, water of

this composition will have no evil effect at 28 days upon the grout with

which it is gaged.

Three series of tests were undertaken, in which a 1:3 mortar of Ot-

tawa sand and a cement of good quality was mixed with Croton water,

and with two typical rock waters encountered in tunnel work.

TABLE 22

Analyses of Rock Waters

Parts per million

E W

COP T 0 RRR RTH ei ee eae ere ea) 85 943

INES Oto Si 2 spe SES ete rea eee cera eeeapeate 159 156

SOs Saar eve oi Sie antatoea os tag atl ncean cobayar ie ans Ree yeep 73 172

IER Ae Sasori pote ear ep oe sae Smee ee URNS 1380 3420

Total solids ..... Pe ral EA ce rae gM Do ge . 2978 7929

The normal amount of water was used to mix the mortars in each

case, and the briquettes were stored in the damp closet over the stated

periods.

TABLE 23

Tensile Strength of 1:3 Mortars Mixed with Various Saline Waters

Pounds per square inch

Number of

Mixed with— — | specimens in

7 days 28 days 3 months phe

Crotomuwaters cn) aoe a eer 302 322 344 6, 5, 6

Water (HS ctu cre Mere Denia 297 343 363 6, 6, 6

Water sob ics tc). et yok Bese 296 335 383 6, 6,6

PAOINI, METAMORPHISM OF PORTLAND CEMENT 213

As was found in the case of the grouts last mentioned, waters of this

general concentration do not appear to affect the strength of cement

mortars with which they are gaged, and the probabilities are that no

serious effects will result from this cause alone.

QUALITY OF WATER THAT MAY SUBSEQUENTLY COME INTO CONTACT

WITH THE SYSTEM

Theoretical considerations —The action of dissolved salts in water

that comes into contact with concrete, where such action is deleterious

to the concrete, has been carefully studied by a large number of investi-

gators (68, 81, 96, 112). Of the salts which have been found injurious,

magnesium sulphate and magnesium chloride seem to have the greatest

effects. What concentration of dissolved salts is necessary in order that

disintegrating effects shall manifest themselves cannot be definitely

stated. This is a field problem and is subject to wide variations under

different conditions.

A water containing relatively little dissolved material, acting under

favorable conditions of porosity, pressure and wide temperature changes

upon one concrete, may accomplish failure of the structure; while

another water, of high saline content, meeting a dense, impervious con-

erete, not forced through the mass by pressure and under conditions of

small temperature change, may have practically no action. Manifestly,

unless these varying conditions are taken into account, it is unscientific

to draw any conclusions regarding the attack of different waters or the

resistivity of different cements.

It may be laid down as a basic principle, however, that the denser a

concrete, other conditions being equal, the greater its resistance to the

attack of saline waters (10, 41, 57). The alkali waters of the Western

states have given a great deal of trouble in concrete construction. Most

experimenters conclude that their action upon concrete is in the main

mechanical and due to the disruptive force of crystallizing or efflorescing

salts deposited in the pores by intermittent submergence and drying

out (30, 38, 49, 56). ;

Of course, as has been pointed out, action of this sort is not confined

to concrete, and any material of construction possessing porosity is

lable to a similar disintegration. The remedy, therefore, is to prevent

the penetration of the saline solutions by the employment of courses of

permanent, impenetrable materials, preferably asphaltic layers.

Where the attack is not mechanical but chemical, this remedy is also

applicable. Unfortunately, there are examples of construction which

are exceptions, and, in these, some change in the chemical or mechanical

214 ANNALS NEW YORK ACADEMY OF SCIENCES

constitution of the cement is the only way to prevent decomposition. In

concrete block construction, where the blocks may be made long before

they are actually put into the structure, it is found of great advantage to

allow them to harden in air or in damp sand, and so permit to a great

extent the carbonation of the lime compounds. Some investigators claim

excellent results from this method (41, 55).

As to the modifications in the constitution of the cement that will

combat the action of saline solutions, there is a great disparity of opin-

ion, which possibly is based upon lack of standardization of experimental

conditions. It is generally conceded that high silica cements are best

suited for the purpose (7). The use of puzzolan cements, or of addi-

tions of puzzolan to the cement in use, is also well recommended (7, 37,

66); and the addition of clay, burnt or dehydrated, finds favor with

some (7,75). As to the lime content of the cement, opinions are divided

whether it should be high (5, 41) or low (92).

Cement of greater density (57) and cement ground to a greater fine-

ness than usual (72) are favorably commented upon. The subject,

because of its great complexity and because of the questionable value of

laboratory results, is at present in a chaotic state. The length of time

that must elapse before judgment may be passed upon the permanence

of a material under these conditions and the corresponding newness of

the field of Portland cement render present conclusions largely a matter

of speculation.

Effect of storage in various saline solutions upon the strength of

mortar.—In order to study the relative resistance to saline solutions

offered by cements varying in chemical composition and in fineness of

grinding, a series of 132 2-inch mortar cubes was made up, in the pro-

pertion of 1:3, with standard Ottawa sand, the cements used being

A. A high silica cement

B. A low silica cement

C. A cement of ordinary composition, sifted and remixed so that

98.8 per cent passed the 100 mesh sieve and 88.6 per cent

passed the 200 mesh sieve

D. The same cement as C sifted so that 92 per cent passed the

100 sieve and 75 per cent passed the 200 sieve

PACINI, METAMORPHISM OF PORTLAND CEMENT 915

TABLE 24

Analyses of the Cements Used in Tests with Saline Solutions

Per cent

A B Cc

SlOke casera ie ie ee as 23.50 19.74 22.99

GLO)! sereye tha let anes 2.36 2.75 2.42

I O)s eats ener asi tees 7.28 8.77 6.79

JEG) se apa aa ee ean Bae 62 18 60.86 60.84

Ce (De Ne ee ee 2.29 2.86 4.14

SOS REESE A aoe ne ean Meelelt 1.39 1.76

CO,H,O, alkalies...... 1.28 3.63 1.06

The cubes were stored 24 hours in the damp closet, and then trans-

ferred to the solutions mentioned in the following table, three cubes to

each liquid, and there stored for three months, then broken.

TABLE 25

Compressive Strength of Mortars Stored for Three Months in Various Saline

Solutions

(Hach value is the average of three determinations)

Pounds per square inch

Storage medium Easy é ' 5

High aa Low ae Finely wey Coarsely a

silica Gani silica ont ground cent | ground een)

Croton water:..| 2217 |...... PAGI. Naa < CARI: 3: ercrete 2066

Sodium 5%| 2267 2 2090 —1d 3266 53 2273 10

sulphate, 10%| 3264 47 2035 —18 2223 4 2262 9

Magnesium 5%| 3244 46 | 1787 | —28 | 2233 5 | 2759 33

sulphate, 10%| 2604 18 | 2646 7 | 3003 42 | 2489 20

Sodium 5%| 2365 7 1785 —28 2305 8 2695 30

chloride, 10%| 1778 —20 2019 —18 2968 40 2044 —1l

Magnesium 5%| 2331 Sale 1827 26) | 273i 33 | 2305 12

chloride, 10%| 1757 —21 1769 —28 | 2570 20 2269 10

Calcium 5%| 2653 19 2516? 2 2219 4 1808 —13

chloride, 10%| 2224 0 1994 —19 2042 —4 2238 8

Average gain — sos,

(percent) 2.4) --.- LOH ele eel eae CG Cie ae 12

9 Average of two determinations.

916 ANNALS NEW YORK ACADEMY OF SCIENCES

The general deductions from these experiments for the period covered

are that the high silica cement, notwithstanding its slower rate of har-

dening, resists the action of these dissolved salts better than the low

silica cement, and the finely ground cement better than the coarsely

ground. Moreover, with the concentrations used, the stronger solutions

in nearly every case had a more destructive effect upon the strength of

the mortar than the weaker. |

The strengths here obtained by storage in salt solutions are in general

decidedly greater than those obtained by storage in fresh water. Hx-

amination of the cubes, when removed from the solutions at the end of

the test period, revealed under a lens that the exterior was being at-

tacked, minute pittings being quite distinct.

The strength attained by these specimens may be considered as a re-

sultant of the balancing of two effects: the deposition of crystallized or

precipitated material in the voids, which by packing the spaces with

solids will increase the compressive strength; the creation of additional

voids by direct solution or by the disruptive effect of metathetically pro-

duced material. It is probable that the disintegrating effect for these

concentrations is reached considerably beyond three months’ exposure.

From the increases in the compressive strength, it is likely that at this

period a great deal of crystallization or precipitation has proceeded,

overbalancing in the main the disruptive effects. This is a general

deduction, and single instances are notable in which the reverse holds

good.

In the case of the finely ground cement, the density of the mortar

made therefrom has prevented the disruptive effect to a greater degree;

and thus the deposition, while not necessarily as much as in the coarser

cement mortars, has had a more marked effect in increasing the strength.

Effect of storage in rock water upon the strength of lean cement

mortars.—A series of briquettes of 1:4 Ottawa sand mortars was made

up, using a normal Portland cement of high quality. The mix was made.

lean purposely to accelerate whatever disintegrating effect might occur.

Batches of the briquettes were stored in bottles in the laboratory for the

7-day and 28-day tests, and additional series were stored in the field, for

the longer tests, at stations where the waters in question were encoun-

tered. The field series were stored in running water, and the action

upon these should be more severe than upon the laboratory specimens

stored in still water. In each case a parallel test was made by storing a

series in pure drinking water. |

PAOINI, METAMORPHISM OF PORTLAND CEMENT 217.

TABLE 26

Tensile Strength of 1:4 Mortars, stored in Rock Water

Strength, pounds per square inch

Water Stored in laboratory Stored in field Secure

7 days| Gain |28days| Gain ||3mos.| Gain |6mos.| Gain

Drinking..| 211 |...... DOT | Meee 27) |e vee 324y ae 12,12,6,6

CO Oo 220 |+4% | 312| 5% || 323] 1% | 247 |—24%| 12,12,6,6

B? )..) 203 |-4% || 287 |=3z% || 318 |—2% | 303 |\— 5%) 12,12 6,6

Ce: OPQ mee 221 |14% | 288 |—3% || 340| 6% | 328| 1%] 12,12,6,6

TABLE 27

Analyses of Rock Waters in Previous Experiments

Parts per million

A B Cc

EES the agit ealas.: 44

SHO), asian ee ee 20 15 4

Fe,0,+Al,0,.......... 7 5 4

CAO Re sae tues 284 399 87

GOR eer a ghee 124 118 38

See nies Ca eck 727 353 31

(OUR rca ted le the moa 826 046 270

CO,, Alkalies, ete...... 949 459 317

Total solids....... 3037 1895 751

The drinking water used to store the blanks contained in neither case

more than 100 parts per million of total solids.

The most consistent reduction of strength, although a slight one, is

observed in the case of water B, a fairly typical sulphato-chloride water

according to Clarke’s classification (18, p. 190). A strikingly high and

sudden reduction occurs at six months in water A, a sulphate water

charged with hydrogen sulphide, while water C, a chloride water, shows

no marked reduction of the strength, which, however, may be due to a’

low salinity.

The six-month briquettes stored in water A showed superficially much

minute pitting, due to the removal of the sand grains, presumably by

solution of the matrix of the cement. Two sections were cut from one

of these briquettes, one transverse and one longitudinal, in the hope of

discovering whether any replacement of the original material by sul-

218 | ANNALS NEW YORK ACADEMY OF SCIENCES

phates or sulphides was going on. The microscopic examination did not

reveal anything of the sort, the sections being in all respects similar to

sections cut from the briquettes stored in drinking water. It was con-

cluded therefore that the loss of strength was due to actual removal of

material by solution rather than by replacement with material which

would cause disintegration through a discrepancy in volume.

The legitimate general deduction from these tests is that, over the

period of experiment, the effect of these waters is greater in void filling

by crystallization or precipitation than in disintegration by solution or

disruption. —

The void-filling material, if of a stable nature and not likely to return

into solution, should be in a measure a protection against the further

entrance of the saline solutions. It has been mentioned that this prop-

erty has been suggested of magnesium hydroxide (70). Probably upon

this possibility is based the reported effect of chemically inert fine ma-

terials, added to the cement for protection against such destructive

action.

SUMMARY OF EXPERIMENTAL RESULTS

1. Increase of temperature of the water with which cement is mixed

causes acceleration of the set up to a certain maximum perenne,

then a retardation.

2. Storage in cold water, without freezing, retards the hardening of

neat cement, and that of mortars more.

3. Increase in the proportion of fine particles in a cement decreases

the permeability of mortar made therefrom.

4. Mechanical agitation increases the strength of cement up to a cer-

tain maximum time; after which, if continued, it reduces it.

5. The setting of cement is accelerated by dryness of the atmosphere.

6. An excess of mixing water progressively reduces the strength of

cement, This effect is partly reversive of itself, and the reversion may

be increased by additional colloidal material in the original cement.

7. Water percolating through concrete dissolves the lime of the ce-

ment chiefly, and this effect tends to neutralize itself by “healing.”

8. Percolation through concrete preferably follows the bedding planes.

9. Salts in solution in the mixing water tend to lower the strength of

cement. This effect may be neutralized by precipitation in the pores.

10. Storage in saline water affects low silica cements more than it

does high silica, and coarsely ground cements more than it does finely

ground cements.

PACINI, METAMORPHISM OF PORTLAND CEMENT 219

GENERAL CONCLUSIONS

In general, the metamorphism of Portland cement represents on an

accelerated scale the processes which occur in natural rocks. The accel-

eration is of course due to the ease with which water has access to the

finely comminuted particles in the initial stages of metamorphism. Many

of the minerals found in natural rocks, when ground as finely as, or finer

than Portland cement, undergo vastly accelerated reactions in the pres-

_ ence of water; colloidal bodies are thereby produced, and the water is

rendered alkaline (18).

The end product of prolonged water action on Portland cement bears.

a striking qualitative similarity to the end product in the kaolinization

of feldspars. The same transformations evidently occur in both cases,—

the alkalies and the lime are abstracted, and the water and alumina con-

tents increased. The exceeding fineness and high adsorptive power of

the resulting products are also similar. The action of water on nearly

all silicate minerals is, in effect, a repetition of this process.

The peculiar adsorptive properties of colloidal bodies render these

hable to coagulation. As has been pointed out in preceding pages, much

of the cementing material of conglomerates and sandstones, except where

calcitic, may have its origin in a similar phenomenon.

On a natural scale, the action of water is greatly retarded, because of

the larger size of the bodies acted upon, and the consequent paucity of

surface upon which water may exert its influence. When Portland ce-

ment has properly undergone its initial metamorphism, the setting

process being complete and the hardening process in great part so, it

approaches the condition of a natural metamorphic rock, and activities

towards its further change are katamorphic and vastly slower in their

results than the initial changes. The component particles have now

become consolidated and the surface offered to the action of water is

minimized. Of course, this is truer of neat cement than mortar and

truer of mortar than of concrete, these being in the order of increasing

porosity.

The hypothesis that crystal formation is responsible for the strength

of hardened cement is not so complete and satisfactory as the colloidal

hypothesis just referred to. In a compact mass, the growth of crystals

can hardly be considered anything but an element of weakness. As has

been shown by the foregoing results, the effects of varying some of the

conditions of the action of water upon cement are best explained by

considering the hardening a coagulative process rather than a process of

crystallization.

999 | ANNALS NEW YORK ACADEMY OF SCIENCES

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lic Cements. (Paper read before Assn. Ger. Port. Cement Mfrs., Ber-

lin, 1907.) Cem. Hng. News, separate.

: The Hardening of Cement Under Water. (Paper read before

Assn. Ger. Port. Cement Mfrs., Berlin, 1909.) Cem. Hng. News, separate.

. MicHae.tis, W. A., Sr.: Puzzuolan Mortars in Sea Water. Tonind. Ztg.,

33, 1308. 1909.

. MoNTEMARTINI, C.: The Hydration of Cements. L’Industria Chimica, 7,

GOD 90S:

. NEWBERRY, S. B.: Cement for Sea Water Construction. Cement Age, 9, 38,

1907.

. NEWBERRY, S. B. and W. B.: Constitution of Hydraulic Cements. J. Soe.

Chem. Ind., 16, 887. 1897.

. O'Hara, J. M.: Action of Sea Water on Portland Cement. Cem. Eng.

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. Prererson, P. M.: Determination of the Finest Powder in Portland Cement.

Tonind. Ztg., 33, 1687. 1909.

. PIERCE, G.: Destructive Action of Alkali upon Cement. Mining Sci., 63,

130. 1911.

. PLtums, R. A.: Waterproofing Concrete. Canadian Cement and Cone. Rev.,

4, 91.

. Porrson, L.: Studies of the Chemical Influence of Sea Water upon Port-

land Cement. Ciment, Nos. 6, 7. 1910.

. Porrer, C. J.: Chemical Changes in Portland Cement Concrete, and the

Action of Sea Water thereon. Jour. Soc. Chem. Ind., 28, 6, 1909.

. Poutsen, A.: Long Time Concrete Tests in Sea Water. Eng. News, 64,

3. 1910.

. Progress Report of the Joint Committee on Concrete and Reinforced Con-

crete of the American Society for Testing Materials.: Proceedings, 1909.

PACINI, METAMORPHISM OF PORTLAND CEMENT 923

. ReaD, EH. J.: The Crystalline Products of the Hardening of Portland

Cement. J. Soc. Chem. Ind., 29, 735. 1910. (Discussion by Blount.)

. REIBLING AND R&yES: Physical and Chemical Properties of Portland

Cement, III. Phillip. Jour. Sci., 6, 207. 1910.

. RICHARDSON, CLIFrFoRD: Portland Cement from a Physico-chemical Stand-

point. Atlantic City, June, 1904. Also, Eng. News, Aug. 11, 1904, Jan.

26, 1905, Cement 1904-1905, 5, 3, et seq.

. Rowan, W. D.: The Effect of Sea Water, Alkali Water, and Sewage on

Portland Cement. Eng. Cont., 24, 52. 1905.

. ROHLAND, P.: The Weathering of Stones and Mortars. Zeits. Chem. Ind.

Wolloide, 8, 48. 1911.

: Influence of Electrolytes on the Setting Time of Cement. Zeits.

Angew. Chemie, 16, 622, 1903. Stahl u. Hisen, 28, 1815 1908.

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ANNALS NEW YORK ACADEMY OF SCIENCES

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of ee and similar matter.

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THE LIBRARIAN,

New York Academy of Sciences,

= care of

American Museum of Natural History,

. New York, N. Y.

- ANNALS OF THE NEW YORK ACADEMY OF SCIENCES

Vol. XXII, pp. 225-258, Pll. XXINI-XXXIIl

Editor, EpMuND Otis HOovEyY '

THE PHYSIOGRAPHY OF THE PERUVIAN

ANDES

py itt NOTES ON EARLY MINING IN PERU

V. F. Marsters

NEW YORK

PUBLISHED BY THE ACADEMY

1% SEPTEMBER, 1912

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Naturau History, 1817-1876) :

OFFICERS, 1912

President—HMERSON McMit.in, 40 Wall Street’

Vice-Presidents—J. EDMUND WoopMAN, FREDERIC A. Lucas

CHARLES LANE Poor, R. 8. WooDwoRTH

Corresponding Secretary—HENryY EH. CRAMPTON, American Museum ~

Recording Secretary—Epmunp Otis Hovey, American Museum

Treasurer—HeEnry L. DoHeErty, 60 Wall Street

Librarian—RatPH W. Towser, American Museum

Editor—EpmunD Otis Hovrey, American Museum

SHCTION OF GHOLOGY AND MINERALOGY

Chairman—J. EH. WoopMan, N. Y. University

Secretary—Cuar.es P. Berkey Columbia University

SECTION OF BIOLOGY

Chairman—FReEpERIc A. Lucas, American Museum

Secretary—WiLL1aAmM K. Gregory, American Museum —

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—CHARLES LANE Poor, Columbia University :

Secretary—F. M. PEDERSEN, College of the City of New York

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. 8. WoopwortH, Columbia University

Secretary—FREDERIC LiyMAN WELLS, Columbia University

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. ACADEMY OF SCIENCES, Vol. XXII," pp. 225-258, Pll. XXIII-

XXNIII. 17 September, 1912]

THE PHYSIOGRAPHY OF THE PERUVIAN ANDES

WITH

NOTES ON EARLY MINING IN PERU

By V. F. Marsters

(Presented in Abstract before the Academy, 5 February, 1912)

CONTENTS

: Page

i TEMTTE DE TO MCULO ING! 5 Bs ceerchersie Ses Sle Gio ee a ae ORC AO LORE ene Ra naar ECR ee ea 225

PO pPOsiAap li GaPEOVINGCES OF REE. a5 Sb oo Sete cieiee ao Sion ie ele dele cele sa Ske dee eats 227

COBDS TAL TOIEMIMSES Se Sos sche co CIS NCR aE nN CECI CR CT a le 228

Orie ME iM aTye MCA rn ac tok situa c cise wie erst aloes w lscvesein sd sc etene ss (20D

Locus of agriculture and its dependence upon physiographic

[DLIVETIVOUINVETTS 653 os ate aro oro G GESTS Han TE RECT er erst eM

Wiest slope and West Range of the Andes........-.....5...-.--+20-- 239

Hira o-Ovone sSeChlOMe sos sobs min cle sicko 's Gil ciate cis ceeds ero doce es coe LOO

@cona-CoOrro, uno? SCChHON sc) kas sic see os a oe bs dls Seale Sees 240

Highland Plateau or Intercordilleran Belt........ PAT eeen or tol eae

WETLOME CMR ASCOU SE CHHOMe i icic\ecrens siete vc es eS bcesele # co's ea ee bese a, ccs, LAD

Casma-Huaraz-Huacaybamba section............. 0. ce ee eee ee 2AG

Piuiraskiunan cabanas SCCHOMs Jac cco vise celiiseis cee cccleocecceescuwes 24T

Hasthanzerand Hast -Slopesc.....2c5.ee2. cece ees ee at Veh ietena Stree 248

Topographic expression as related to the geology of the Peruvian Andes. . 249

Pal eMUM Sp cUIN ESOC IT OIMGS are sta crane ene el cue cca .c) eiisie a's couch aveve, ace we 14) ots teeta. 6 Steeles: 26 249

ZoOrritos-Wambeyagque Plait. . ose. 0 cote ss oe Sas cane cece cee eevee JAD

Ghaimehra-O lino Ss Ane seer ares gins: uayens ate ste Sieae-ce ees Seow oie ieee =.) LOO

@cona-Moquequa” S@ChHON. 2.4.0. cc cc. secs ces croc een es cer ees sees, ZO

Huacho-Cerro de Pasco section................. tins ei areoeeeseaaiccoin ee 251

WMeona-COLeoe UN OMSECH OME dem elocra ces cies Hele A cele oe et BAS ee shee os OD

NL CSROM CARLY aIMIMIN SINE POR IWs seals nies ee ois = Selelelo ses isle ebles viene ase) 2D

Ee UIA ecgets, eWay nersno loca eres ets goeani's Gy Seah nits, Gata re aC mAela ws. 8 AAs ae atksicceniyere EOS

INTRODUCTION

So far as I am aware, very little has been written relative to the

physiography either of the coast or of the Peruvian Cordillera. As

preliminary to the presentation of a few observations on the above

subject, I wish to summarize very briefly a list of the early pioneer

geographers, geologists and naturalists who have visited the Peruvian

shores and have published observations relative to the subject in hand.

(225)

226 ANNALS NEW YORK ACADEMY OF SCIENCES

Darwin was among the first to record observations relative to the

coast of Peru. He noted the many old shore line beaches now standing

at various elevations above the present sea level. The same facts were

noted later by D’Orbigny. The occurrence of “kitchen middens” was

recognized at various points. They appeared to be associated with the

ancient beaches, thus suggesting that elevation had actually taken place

since the accumulation of the shell heaps. The writer has seen between

the mouth of the Rio Grande and Lomas as many as five well-defined old

shore lines or beaches occurring in succession, the highest one being

approximately some sixty feet above sea level.

I find also that a geographical map of a portion of southern Pern,

together with a portion of Bolivia and Chile, has been prepared by one

Mr. Pentland, but to date I have been unable to locate it or any of his

written contributions on the geography of the above section.

Pissis and D’Orbigny were the first to contribute detailed geological

and indirectly geographical information concerning southern Peru and

the adjacent republics. Both these men constructed cross sections of the

Peruvian and Bolivian Andes, the section starting at the coast near Tacna

and ending on the east slope of the East Cordillera to the east of La Paz.

Some years later Forbes went over the same ground and likewise con-

structed a cross section. While the same type of relief is expressed in all

these sections, the classification of the formations, based in part upon

lithological and in part upon paleontological data, differs very widely in

each case. The sections prepared by these men may be seen in the now

rare publication of the Geographical Society of La Paz, Bolivia.

Among the later pioneer naturalists who did much serious work in

Peru was one Senor Raimondi, an Italian by birth. He came to Peru,

bringing with him the training of an Italian institution and that pro-

found interest in his field of investigation that is always sure to produce

invaluable results. Raimondi was the Agassiz of Peru. Among his

first efforts was the preparation of a topographic map of Peru; and to

date it is the only map possessed by the republic and officially accepted

by the government. The writings of Raimondi are likewise voluminous.

While collecting his map data he likewise accumulated a mass of infor-

mation relative to the geology, mineralogy, zodlogy and botany of the

entire country. The results of his investigations were published in a

series of volumes by the Peruvian government.

Another source of information is to be found in the publications of

the Cuerpo de Ingenieros de Minas, a department of the government in-

stituted by Sr. José Balta for collecting information concerning the

natural resources of the republic, such as mines and mine production, the

Nt rn pi mines ody

eee SoH Aipeme We ven wae pn Sed ee ete wy Sey ee eae Lo

ee nT a ET UREN Ca acer aT S

ANNALS N. Y. Acap. Screncps

CUAYRQUIL a iT

NESS.

{

Lobosise, Ete, 7,

ESI

Trujillo i

CALLAO KE”

SKETCH MAP

SHOWING THE

GEOGRAPHICAL PROVINCES OF PERU

V. F. MARSTERS

FS] Coastal Ridges

Coastal Plains

West Slope-W. Cordillera

East and West Cordillera

Intercordilleran Belt

a, %

TOP SBUGAIACES Of

CEOCE YEH 0 :

cee settee

a erenepersabaeehir ote

7 ST

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 99%

geology of mining districts, data concerning irrigation projects, under-

ground waters, etc. More than two hundred bulletins have been pub-

lished by this department. While much information exists in the series

of publications, it is unfortunate that no attempt has ever been made to

correlate it with a view to preparing atleast a preliminary geological map

of the more important mining centers of the republic. So far as I am

aware, the only attempt to correlate the formations in distinct parts of

the republic was recently made by Dr. G. Steinman for the Cordillera an:]

by the writer for the formations of the coastal plains as they appear in

northern, central and southern Peru. Hence we may say that, while there

are sources of information relative to the geology and geography of the

Peruvian Cordillera, in no publication, so far as I am aware, has any

attempt been made to give even a skeleton outline of the probable physic-

graphic history of any section of the Andes.

Let us now see if, from the information to be gleaned from the early

writers and especially the map of Raimondi, combined with the observa-

tions of the writer, we may be able to get a concise picture of the

geography and at least a glimpse into the geology of a portion of the

Peruvian Andes.

The Republic of Peru occupies an area approaching 2,000,000 square

kilometers ; it extends from 0° to 20° 8. latitude and from 64° to 84° W.

longitude; its coast line is approximately 1,500 miles long, and it pos-

sesses all the varieties of climate from typically tropical conditions in the

north to cold temperate in the south and in the higher parts of the

Andean Highlands. One need not go beyond the confines of the Inca

Republic to find any of the variations between the extremes mentioned.

One may leave Lima under a semi-tropical sun and, in the course of a few

hours by rail, be riding over a snow-covered Puno. As soon as he reaches

the montana, or the wooded part of the eastern slope of the east range, he

passes immediately into warm temperate and tropical climates.

TopoGRAPHIC ProviINcres oF PERU

The distinct topographic and physiographic provinces of Peru are well

defined. They may be summarized as follows:

1. Coastal Plains.

2. West Slope and West Range of the Andes.

3. Highland Plateau, or Intercordilleran Belt, and its associated sec-

ondary ranges.

4, Hastern Range.

5. Kastern Slope and Lowlands.

298 ANNALS NEW YORK ACADEMY OF SCIENCES

COASTAL PLAINS

The distribution of the coastal plains may best be seen in the accom-

panying sketch map. They occupy three sections, which may be designated

as the northern, central and southern divisions. .

The northern division begins in the region of Tumbes, near the bound-

ary of Peru and Ecuador. It extends along the coast to the south, reach- -

ing its maximum width in the cross section between Cerro del Yllesca

and Salitral. From this point to the south, it grows narrow quite rapidly

and finally ends at Salaverry.

Going still farther south, we find but few remnants of what must have

been the inner edge of the coastal plain tucked away in partly drowned

valleys within the limits of the western slope. Between these valleys,

the formations composing the foothills of the western range now occupy —

the present shore line and continue to do so, until we reach the region of

Canete. Here the coastal plain again makes its appearance. It can be

traced to a point just to the south of the mouth of the River Yauca.

Here again the foothills of the western slope reach the present shore line.

If we were to travel along the shore line from Yauca to the south, we

should be very much inclined to believe that, as far south as Ocona, the

actual foot-hills of the Cordillera again formed the present shore line.

This, however, we find not to be true. Should we pass up the Valley of

the Atico, we should find a belt of country some fifteen to twenty miles

wide occupied by Tertiary and post-Tertiary deposits, similar to those we

have seen to the north but standing at a higher altitude. They are deeply

cut by the streams coming from the west range of the Andes. It is not

improbable that this belt may extend to and beyond the Valley of the

Chaparra. Going to the south, we can trace the elevated coastal plain,

where it is known under the various names of Paco Alto, Cuno-cuno,

south of the Valley of Ocofa, Pampa de la Joya, south of the Valley of

the Vitor, Pampa de Clemesi, south of the Tambo Valley, etc., and thus

continues with decreasing altitude to the southern boundary of Peru and

Chile. :

One of the additional features of the coastal plain is the frequent

occurrence of isolated hills and ridges near the present shore line. These

are locally known as “morros.” They usually stand-at an appreciable

elevation above the general level of the plain and hence form convenient

landmarks for determining direction of travel to a given point.

With the exception of the section of the coastal plain to the north of

the Valley of the Ocofa, the principle transverse streams have succeeded

in cutting only deep narrow valleys, with very little available floor for

“MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 229

agricultural purposes. The interstream spaces are barely scarred by insig-

nificant tributaries, so that we have huge expanses of desert plain extend-

ing from valley to valley. As we approach Moquequa and Locumba, how-

ever, the elevation has been less, so that the rivers have made broader

valley floors on which has been developed considerable grape culture,

while, in the Valley of Sama, sugar cane forms the chief agricultural

product.

Geologically considered, the coastal plains of Peru are composed of

Tertiary and post-Tertiary sediments and lava flows, laid down on a

post-Cretaceous surface, which at the beginning of Tertiary sedimenta-

tion had not been worn down to grade. This is proved by the number of

half-buried hills now standing above the general horizon of the coastal

plain, the so-called “morros” already referred to. In many instances,

these hills merge into ridges and form what is known in Peruvian

geography as “Cadena de la Costa,” or coastal chain. Their distribution

may be seen on the accompanying sketch map.

As might be expected of a coastal plain bordering a mountain range

of such tremendous proportions and geologically young, it has under-

gone differential elevation to a great degree. In various localities, folding

and faulting have taken place on a considerable scale.

~ In that section to the-north of Paita where I have made some detailed

stratigraphical studies, on account of the development of the Zorritos,

Lobitos and Nigritos oil fields, it is caleulated that not less than 3,009

feet of Tertiary sediments enter into the structure of the coastal plain.

Some localities are rich in fossil gastropods and nautiloid forms. They

are supposed to represent lower and middle Tertiary faunas.

It is interesting to note the fact that the oil-bearing localities so far

developed are associated with the sections of maximum disturbance of

the formations. The oil field of Zorritos is located on the eastern flank

of a folded section, much of which is located beyond the present shore

line. It is also in this section that we find the largest amount and most

minute type of surface dissection. The folding of the Zorritos section

may be traced to the northeast for some distance. In the interior to the

southeast, the folding gradually fades out until we reach the flanks of

Amotape Mountain, where its contact with pre-Tertiary formations is

encountered.

Passing to the section of Lobitos, we find the same stratigraphical and

structural relationships as noted in Zorritos. The area of maximum fold-

ing is near the present shore line. About Lobitos, the original surface

of the coastal plain has been completely eroded away. To the north of

the Lobitos field, we find the original surface forming an extensive plain

930 ANNALS NEW YORK ACADEMY OF SCIENCES

some 300 feet above sea level and with its shoreward edge somewhat cvt

up by narrow, short valleys. Passing to the interior from Lobitos in the

direction of the Amotape spur of the west range of the Andes, consid-

ered in its broadest sense, the evidence of folding, faulting, etc., rapidly

disappears, until we approach the inner edge of the plain, 2, we find

that the Tertiary sediments are upturned.

Following to the south towards Talara and Nigritos we find that while

folding is much less accentuated than at Lobitos, it by no means disap-

pears entirely. At Talara, however, we find another area of maximum

disturbance, which extends to the southern end of the oil field now being

operated by the London Pacific Petroleum Company. Following this

section to the interior, we note, as in the preceding cases, a disappear-

ance of the folding, until we reach a narrow belt adjacent to the Amotape

Mountain, where the Tertiary strata are found to stand at a high angle

and dip toward the northwest. I wish here to record another fact. The

first oil springs noted in this region were located at the interior edge of

the coastal plain at a point known as La Brea. A little later a residuum

of petroleum was located on the playa near Nigritos. This was mined for

some time by the Spaniards and prepared for painting the bottoms of

their ships.

In the Chira Valley, we find the Tertiary occupying an arm-like ex-

tension or depression between the Amotape spur on the northwest and

an edge of the foothills on the southeast. On the high plains to the rear

of Paita, some 300 feet above sea level, there is a well-defined noncon-

formity between red clays with a marked tilt and a series of sands and

conglomerates containing numerous fossils. Many of these resemble very

closely the forms now living on the present shore. At various points

between Paita and Piura there may be seen on the surface shells of

exactly the same variety as are now living on the Pacific shore.

Just south of Paita we find a typical outlier of the western range stick-

ing its higher points somewhat above the level of the adjacent plain.

In the interior, the eastern boundary is located near Tambo Grande. It

follows the general trend of the rivers Salitral and Serran (or R. Piura),

or not far from the contact of the coastal plain deposits with the pre-

Tertiary formations of the foothills. That is to say, the Salitral-Serran

valley has been made by a longitudinal subsequent river, using the phrase-

ology of Professor Davis.

The maximum width of this division of the coastal plain is attained

along a line from Cerro de Yllesca to Salitral. Following the inner edge,

we pass to the south through Olmos, Motupe, Patope and Cayalti. A very

narrow belt then passes along the present shore line to the Pacasmayo

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 931

Valley. The Cerro de Yllesca forms another of those outliers that serve

to break the monotony of the sky line as seen from the interior.

From the Tablazo de Paita, the elevation decreases somewhat in the

direction of the Despoblado de Sechura and the Plain of Olmos, while a

minimum uplift took place in the section through Morrupe, Chiclayo

and Lambeyaque. As to the stratigraphy of the Sechura-Olmos area we

ean say but little. This is one vast plain, a typical desert, strewn with

wind-blown sands, with here and there small depressions occupied with

“salinas” or salt deposits, but so slightly dissected that data giving a

clue to the stratigraphy are wanting. Sections, however, between the

mouth of the Sechura River and Cerro del Yllesca, as exposed on the

shore line, lead me to mistrust that beneath the sheet of recent accumula-

tions over this vast plain there must exist a thick series of Tertiary sedi-

ments. Not a single morro of pre-Tertiary origin is to be found until

we approach the region of Olmos. The accumulation of Tertiary sedi-

ments in the central parts of the desert may thus be very considerable.

As has been stated, the continuation of the coastal plain below Cayalti

is represented by only a narrow rim, which, as we approach the Valley

of Pacasmayo (the River Jequetepeque), again widens out to several kilo-

meters. Here we find that the outhers of the foothills are quite numer-

ous. An interpretation of the stratigraphy in this valley is difficult on

account of the lack of good exposures. Sufficient evidence is at hand

to suggest that the red clays corresponding with the lower part of the

Paita section are present.

Following to the south again, but a narrow band of the plain bordering

the shore line connects with the Valley of the Chicama and Santa Cata-

lina. ‘These valleys are probably filled in part with late Tertiary sedi-

ments, but later were littered up with much post-Tertiary waste, a phase

of coastal geography to be discussed later. The southern extremity of

the northern division ends at Salaverry.

The stretch of coast from Salaverry south to a point just south of

Lima is largely occupied and confronted by the foothills of the Cordil-

lera. Only the larger valleys have broad floors near the coast. Whether

many of these contain remnants of Tertiary formations or not has not

been determined by the writer. It is not improbable that during Tertiary

time a fiord-like arm of the Tertiary Sea may have extended into the

partially drowned valleys of the West Range. At all events, it is certain

that a large amount of waste has very recently accumulated and spread

over the lower stretches near the present shore line, presumably in post-

Tertiary time. The valleys of Huacho and Chimbote are fair examples.

Mention should also be made of the fact that, in the upper portions of

232 ANNALS NEW YORK AVADEMY OF SCIENCES

many of these partially filled pre-Tertiary valleys the recent deposits of

sand, gravel and clay have again been attacked by the present streams and

redistributed at lower levels. The same type of physiographic history is

repeated in the valleys of the Rimac and the Chillon. The lower part of

the Rimac Valley and the broad area where it is confluent with the

Chillon have been aggraded by the deposition of an enormous sheet of

fluviatile material. Lima, the capital of the republic, stands on the edge

of this fluviatile plain. Whether Tertiary formations exist beneath the

sheet of waste or not has not yet been determined.

Not until we reach the vicinity of Cerro Azul, Canete and Chincha

does the typical coastal plain again make its appearance. In the section

through Chincha it has a width of some three or four kilometers. ‘The

transverse streams have incised themselves but slightly into the surface

deposits. The formations thus far exposed here appear to be post-Ter-

tiary waste, sands and conglomerates, undoubtedly deposited upon late

Tertiary clays. The latter may be seen in sections at points between

Chincha and Pisco to the south. These are regarded as the equivalents

of the late Tertiary clays at the bottom of the Paita section. Folding

and minute faulting may be seen at various points; the best exposures

occur near Pisco. To the north of Pisco, the amount of Tertiary deposi-

tion may have been very considerable, but to the south, there is reason to

believe that no great proportions were reached. ‘The fact that a very

large part of the shoreward area from Paracas to the mouth of the River

Ica is occupied by outliers of the pre-Tertiary oldland, precludes, at least,

the idea of Tertiary deposition on any great scale in that particular sec-

tion. It was only between this broken line of hills or outliers and foot-

hills of the west range that any great amount of Tertiary /leposits accu-

mulated. In the upper part of the Ica Valley, within the limits of the

coastal plain, a very thick sheet of post-Tertiary waste was spread out on

the Ica plains as far south as Ocacaje. To the west of Ocacaje, the best

exposures of the light-colored Tertiary clays containing fossil fish are

to be seen. These rest upon the eastern flank of the outhers already

referred to and probably pass under the fluviatile plains in the upper

stretches of the Ica Valley and plains. From Ocacaje south we may trace

the plain across the Pampa de Huayuri to Palpa, San José and Nasea.

To the south of the Rio Grande, we reach the Pampas de Yunga Colo-

rado and Bella Union on the northwest side of the Valley of Acari. The

plain finally ends at a point a short distance south of the mouth of the

River Yauca. The Pampas de Yunga are separated from the coast line

by Cerro Yunga, composed of pre-Tertiary formations, another outlier of

the West Range. Some distance to the south of Cerro Yunga, the coastal

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 233

deposits again form the shore line and continue with but little inter-

ruption to the southern terminus of the central division. From the

Pampas de Yunga, with an elevation of some 700 to 900 feet above sea

level, the surface gradually descends, until, at Lomas, it passes beneath

sea level. To the interior from Lomas, where the plain is known as Bella

Union, the surface is some 300 or more feet above sea level. In the valley

of the Rio Grande, sections of what appear to be late Tertiary may be

‘seen. On the coast near Santa Ana, similar sections may be found. On

the southwest flanks of Cerro Yunga, the same sediments are exposed.

In the interior, on the inner edge of the Pampa de Yunga, is the so-called

Vallé de Carbonaria, where a considerable thickness of clays and sands,

presumably late Tertiary in age, may be seen.

To the south of the plains bordering the Valley of Yauca, the foot-

hills once more form the present shore line. Nevertheless, in the mouth

of the Valley of Atiquipa and again in the Chala may be found a tri-

angular patch of what appear to be late Tertiary clays and sands: but

beyond this point the foothills seem to present a solid front, until we

reach the mouth of the Ocona Valley. This, however does not prove to

be the case. Should we enter the mouth of the Atico Valley and journey

in the direction of Caraveli, we should find that the supposed foothills of

west range of the Andes prove to be outliers, separated from the main

range by much-dissected plains made up of Tertiary and post-Tertiary

formations, and that these extend to the actual foothills of the Andes

in the vicinity of Caraveli. These plains probably pass to the northwest

as far as the valley of the Chaparra and represent the northwestern limit

of the south division of the coastal plains. The inner edge of this plain

is not less than 3,000 feet above sea level and has a width of twenty-five

miles or more. The pre-Tertiary coastal ridge as far as Oconia shows only

its highest point at elevations exceeding the general level of the interior

plain, while its shoreward flank has been nearly stripped of its post-Cre-

taceous covering. It is in sections on the Caraveli and tributaries to the

Atico that we find for the first time that mud flows of no mean propor-

tions enter into the formation of the coastal plain. In Quebrada Chin-

chin, we may see at least 300 feet of this flow exposed. Moreover, these

flows extended far in the direction of the coastal ridge. From the region

of the Atico section, we may trace the high plains to the southeast, being

represented by the Pampas de Paca Alta, Bourbon, etc., to the valley of

the Ocona; then we find the continuation in Pampas de Cuno-cuno,

Majas and Vitor, where we reach the valleys of Siguas and Vitor. The

inner edges of these pampas stand at least 5,000 feet above sea level.

They are very deeply cut by the main streams coming from the West

934 ANNALS NEW YORK ACADEMY OF SCIENCES

Range, in fact most of them have cut veritable canyons and, in many

parts, are still continuing the process of incision. The inter-stream

spaces are, as yet, barely scratched by tributaries to the master lines of

drainage. The pampas thus far named are but high-lying plains, with

their initial surfaces well preserved. They are veritable deserts.

To the southeast of the Vitor Valley we find the pampas of La Joya

and Chachendo, the two being partly separated by outliers of the West

Range, while nearer the coast and on the northwest side of the valley

of the Tambo we find the pampas of Islay and Tambo.

Crossing the Tambo Valley, we meet the lower plains adjacent to the

shore line and usually known as the La Punta Plains. Crossing the

coastal ridge, however, we come upon a higher-lying plain of enormous

dimensions and usually known under the name of Clemesi. It extends

with little interruption to the valley of the Moquequa. It decreases very

materially in elevation as we pass from the Tambo to the Moquequa

Valley. On the Moquequa, the plains have a width of from twenty to

twenty-five miles.

Of all the pampas so far mentioned in the southern coastal plain, the

lava flows already mentioned form no inconsiderable part of the deposits

making the Siguas-Vitor La Joya sections. Excellent sections may be

seen on the Siguas and Vitor valleys. Here can be seen two distinct

flows, as indicated by the inter-stratification with sediments. The max-

imum development of the flows seems to have taken place along a line

or section from Arequipa to Quilea, the former town being located well

within the west slope of the West Range of the Andes, and on the con-

tact,of the flow with older formations.

Just what may be the correct correlation of the sediments of these

sections with those of the central and northern division of the coastal

plain has not been determined with any degree of certainty; it is mis-

trusted, however, that they represent rather late phases of Tertiary de-

position. It is important to note, as well, what an enormous amount of

elevation has taken place, since their deposition, simce we now find some

of these at least 1,000 feet above sea level. There is no proof that the

lava plains as a whole ever stood at sea level. The fact, however, that

salt deposits are known to occur well up to the higher plains indicates

a depression of such extent that a part at least of the mud-flow must have

been near sea level. .

Returning to our description of the areal extent of the southern divi-

sion, we find that the Pampa de Clemisi has suffered but slight erosion.

Near the inner edge of the Cadena de la Costa are some very fine salt

deposits. The best sections are to be found in the slopes of the Moque-

PLATE XXIV

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ANNALS N. Y. ACAD. SCIENCES

VoLUME XXII, Phare XXVI

PLATE XXVI

Gena ana ae a pe OELENDO

, { Siw

“sands IS ‘and bays of the coastal plain,

eer ; W ee i ty, \

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Fi IME XX tp XNXV

ANNALS N. Y. ACAD. SCIENCES VOLUME XNII, Phare XXVII

Riaime

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PLATE XXVII

MOQUEQUA (ILO) RIDGE

View of the Moquequa (the Ilo) near the coast where it has eut a deep

gorge in the Coastal Ridge.

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 935

qua Valley and in the Cuesta de Mato de Caballo. Here is éxposed a

series of somewhat coarse deposits on top, followed by a thick series of

clays and fine sands. Folding and faulting have taken place to a con-

siderable degree.

The Pampa de Clemesi has an elevation near the Tambo Valley of

some 4,000 feet on its inner edge, but it descends in the direction of

Moquequa and Locumba. At the valley of Sama, which at present con-

stitutes the dividing line between Chile and Peru, the elevation at the

inner edge is not more than 800 to 1,000 feet. The same grouping of

sediments may be seen on the Sama as in the Moquequa Valley.

As regards the character and distribution of the outliers,.a word must

be added. The pre-Tertiary coastal chain is persistently present from the

valley of Atico to the mouth of the valley of Sama. Near the valley of

the Tambo, the edge of the foothills practically joins the Cadena de la

Costa. It is for this reason that we have a number of small pampas

separated by a collection of outliers. In the stretch of coast from Moi-

lendo to Quilea and Camana, the coastal ridge is most prominent in its

topography and reaches a width of some kilometers. Their topographical

strength, however, gradually fades away in the direction of Sama. To

the south of this point, I am told, the ridge is replaced by a series of

morros, of which the historic morro at Arica is a good example.

Original Tertiary Area—At the present time we probably see but a

small portion of the original area of deposition of Tertiary time. The

Bay of Guayaquil is very shallow and may represent a slightly depressed

part of the Zorritos-Tumbes plains. That Tertiary deposits extend some

distance to sea in front of the Zorritos-Nigritos shore line is more than

probable. This conclusion is borne out by the fact that, for many years

past, mariners have repeatedly reported the occurrence of oil patches on

the surface of the sea some miles out from the present shore line.

Further, such reports are not confined to this part of the coastal shelf,

for quite recently similar evidence has been noted in the region of the

Lobos Islands, located in front of Pacasmayo. Jn fact, so marked was the

evidence that parties of Lima have recently attempted to take up territory

on the above-mentioned islands, with a view to developing a petroleum

industry.

So far as is known to the writer, petroleum-bearing formations of the

coast are confined to the Tertiary horizons. Even in the interior of Peru,

in the region of Lake Titicaca, there is every reason to believe that the

petroleum deposits of the Pusi section are likewise confined to the Ter-

tiary.

236 ANNALS NEW YORK ACADEMY OF SCIENCES

Locus of Agriculture and tts Dependency upon Physiographic Phe-

nomena.—tIn these days, much is said about physiographic features as

determinative factors in the location of human industries. A word con-

cerning the distribution of agriculture on the coastal plain is worth

while.

Mention has already been made of the differential elevation which the

coastal plains of Peru have suffered. Where the elevation has reached its

maximum, or where the plain has been aggraded to a much greater ele-

vation, as by the addition of lava flows, the present streams coming from

the flanks of the oldland (the west slope of the Andes) have cut out verit-

able canyons.. Time enough has not yet elapsed to permit these streams to

widen their floors to any appreciable extent. Here and there in some ot

the canyons, the stream has reached the underlying oldland and its rate

of erosion has been retarded. Above this point, the stream has thus had

some opportunity to do some side cutting and hence has widened its floor ;

only at these points do we see man availing himself of the agricultural

opportunities offered him. In proportion as the elevation has been less

so in a general way, we may say that transverse master streams have had

a chance to make floor room for the use of man. Let us take a few ex-

amples to illustrate the principal stages and their physiographic bearing

on the topic in hand. Probably the maximum elevation of the coastal

plains of Peru was attained in the Mollendo-Arequipa, or Vitor, section.

While formations entering into the structure of this section are in part

sedimentary in origin, it is not assumed that the present elevation of its

surface above sea level represents the actual uplift above that plane. lt

is very probable that this part of the plain has actually been aggraded

by a lava flow, or flows, the present surface of which never stood at sea

level, although the lower part of the earlier flows may have reached the

edge of the Tertiary sea. At all events, the Chile-Vitor stream coming

from the high plateaus to the interior have, within the limits of the

coastal plain, sueceeded in cutting a very deep canyon. Further, it has

developed its widest floor near the contact of the inner edge of the

plain with the pre-Tertiary oldland, or the foothills of the West Range

of the Andes. Fer that reason we have Quercotilla, an agricultural com-

munity and a village, located at this point. Within the remainder of this

valley to the sea, we do not find a single town until we reach the present

shore line, where Quilca, a small village, is not placed on the valley floor,

but to one side in a deep-water embayment of fiord-like character. This

little village is here for the dispatch by water of agricultural products

from the valleys of Camana and Ocofia, neither of which possesses a safe

port of entrance. Mention has also been made of the special case of

AND THE FOOT NGE OF THE ANDES

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XIXX GiVIid ‘IIXX DNOAIOA SMONHIOS “GVOV “X “N STVNNV

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 937

Arequipa. Arequipa is some miles within the interior of the West Range

of the Andes. It stands on the slope and near the edge of the lava flow,

or flows, playing so important a role in the lithological content of the

coastal plain to the southwest. To the rear of the city, and forming a

most picturesque background, the majestic cones of Misti, Pichu-pichu

and other voleanic ridges rear their heads to 19,000 feet or more above

sea level, and 9,000 to 10,000 feet above the light-colored lava plains of

the Arequipa campo. The Chile River coming from the high plateau to

the interior passes between two of these cones, where it has cut a most

picturesque canyon, pushing its way across a part of the mud flow until

it reaches the contact with pre-Tertiary formations of the foothills. The

important fact to note here is that where the Chile succeeded in making

a valley-floor, namely, at the moment it reached the contact referred to

above, there was located the city of Arequipa. Here we find agricultural

pursuits well developed. In all of the southern part of Peru there is no

prettier bit of landscape to be found than the campo of Arequipa. It is

the Switzerland of Peru. That is to say, physiographic features were the

determinative factors in the location of this important trade center.

In the Valley of Majas, we again find that a valley-floor has been

developed near the contact of' the inner edge of the coastal plain with the

edge of the foothills of the West Range. Here has developed a large grape

culture and the manufacture of wine; but from this locality across the

plain we find little chance for the farmer. Not until the Majas passes

through a deep, narrow gorge cut in the coastal ridge, or cadena de la

costa, do we find an additional opportunity for the development of agri-

culture. As soon as the Majas passes beyond the west slope of the ridge,

it has greatly widened its valley, having cleaned away the sediments on

the west flank and built for itself a broad, semi-triangular fluviatile

plain, which has pushed seaward sufficiently to be a menace to the mari-

ner. On this plain, we find the agricultural town of Camana located

and a campo alive with agricultural activities.

The plain on which stands the city of Lima and its suburban towns is

but a repetition of the same physiographic sequences, the difference being

that the Lima plain was built at the edge of the cordillera or foothills

proper of the West Range of the Andes, while the latter developed on the

seaward edge of an outlier of the same physiographic province.

In the Valley of Pacasmayo, we find that the river cut its way to the

sea and at the same time widened its floor throughout its entire course;

as a consequence, no inconsiderable part of the entire floor from the edge

of the foothills to the present coast line is under cultivation, or has been

at various periods. ‘The amount of cultivation is, in some cases, de-

238 ANNALS NEW YORK ACADEMY OF SCIENCES

pendent upon the amount of available water. In the Pacasmayo case we

find an additional feature worthy of mention. At the contact of the

coastal plain with the foothills, the valley has not been deeply incised.

On both sides of the valley are extensive plains, composed largely of the

waste from the edge-of the foothills, plus the original surface of the

coastal plain. So near are these plains to the level of the river, but a

short distance within the foothills, that water has been diverted from its

legitimate place on the valley-floor to irrigate large stretches of plain on

either side of the valley. Success in this attempt has been foiled, in part,

by the lack of water for the extent of territory taken up on the one hand

and the strong tendency to salinity of large tracts of the plains on the

other.

It is only where the coastal plain has undergone the minimum amount

of elevation that we find the last distinct stage to be described. This is

fairly well illustrated by the Etén-Lambayeque-Motupe plains, where we

have a group of small streams coming from the interior to a slightly ele-

vated plain. Only along the inner contact of the coastal plam with the

foothills have these streams slightly incised themselves, but as they ex-

tended outward and over the plain they actually spread their waste over

large areas. Under such physiographical relations, we have the condi-

tions for the development of the most important rice industry in the

entire Republic of Peru. The rice fields occupy the fluviatilly aggraded

portions and such adjacent parts of the original plain as may be reached

by the amount of water available. The growth of sugar cane has also

become an important industry. Where the main streams have formed a

well-defined valley-floor, the predominant culture is of maize, alfalfa and

the staple vegetable products for the markets of the principal towns.

In the valley of the Piura, we find a slightly incised valley, the floor of

which is occupied by cotton culture, as the foremost industry, throughout

its upper stretches. Notable lack of water has limited the territory under

cultivation. Piura and Catacaos are located at these points. Again we

find this portion of the valley-floor near the inner edge of the coastal

plain. The lower portion of the Piura Valley has spread itself over the

plain. Not enough water reaches this part to assure crops. The town

of Sechura is located near the mouth of this sand-laden water-way. The

people here can maintain themselves only in part by agricultural work.

Not a small part of the inhabitants is engaged in transportation, fishing

and salt-mining.

ANNALS N. Y. ACAD. SCIENCES VOLUME XXII, Puarns XXX

Wie. 2

ry ie ait OM

seh eneenne “MOOT Rp so:

Hei. $s sttogts wa ‘aso alt nd OM

ak rae anche Pony

a EE aa t A

aa “ {

Hebkibire’: heal uk Fs: ty twain

Pana f ot, tho ecb ants low Dain Asn) auld, Ho) MG

PLATH XXX

WEST CORDILLERA NEAR CERRO DE PASCU

Fic. 1.—Limestone summits in the West Cordillera as seen in the Oyon-

Cerro de Pasco Section.

WEST CORDILLERA NEAR UTOTO

Vie. 2.—The West Cordillera in the region of Utoto, showing limestones

on the left and voleanics on the right. Ore bodies exist near the contact.

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 229

WEST SLOPE AND WEST RANGE OF THE ANDES

We now come to the geographical province which I have designated as

the West Slope and West Range or Cordillera of the Andes. So far as

I am aware, no one has attempted to outline the physiographic history of

this section, nor is it my intention to try to solve this problem in all its

details ; nevertheless, I wish to present a few of the larger physiographic

features and their variations as observed in various sections on the West

Slope, with the hope at least that these may lead in the direction of a

correct and final analysis.

As a basis upon which to formulate our views, I shall describe what

to me appear to be two fairly typical cross sections of the West Slope in

central and southern Peru. These IJ shall designate as the Huacho-Oyon

section and the Ocofa-Cora Puno section.

Huacho-Oyon Section.—The town of Huacho les about sixty miles to

the northwest of Callao, on the coast, in the valley of the Huari. You

may ascend from Huacho up to the Huari Valley and its important

branches to Oyon. Oyon stands at the base of the more prominent peaks

of the West Range, hence the choice of the two names to locate the section

to be described. The distance from Huacho to the east side of the West

Range is approximately 35 leagues, 175 kilometers, or 105 miles. Begin-

ning at Huacho, we find that the valley of the Huari has a broad flat

floor at the present shore line. It extends inland some eighteen to twenty

miles. It is the seat of important agricultural industries. At the apex

of the Huari floor we find the river running in a deeply incised valley.

Its tributaries have likewise cut canyon-like side valleys. On either side

of the Huari Valley, at the coast line, we find that the foothills of the

West Range come practically to the present coast line. Should we ascend

to a point in the foothills from which a long-distance view of the upland

surface may be clearly seen, we shall at once note a moderately even but

highly inclined surface descending in the direction of the sea and ascend-

ing in the direction of the culminating points of the West Range. As we

pass in the direction of the snow-capped prominences to the northeast,

the inclined surface gives way to prominences standing out in clear relief

above the upland surface. These finally culminate in the peaks of the

West Range of the Andes. In the Lima-Oroya section, the above physi-

ographic features can be discerned, but they are by no means so clearly

defined. If we look into the details of topographic expression on the

West Slope, we find that it has been most minutely cut by steep, narrow

valleys leading to the master transverse lines of drainage from the in-

terior. The formations entering into the structure of the West Slope are

240 ANNALS NEW YORK ACADEMY OF SCIENCES

Cretaceous or older sediments and volcanics. The latest formations, at

least, are not younger than Cretaceous. We also have good reason to

believe that the oldest formations entering into the coastal plain are not

older than early Tertiary. It therefore follows that the topography of

the West Slope must have been developed in post-Cretaceous time, or

more accurately, between the period of uplift of the Cretaceous sediments

and their associated volcanics and that of the initiation of Tertiary sedi-

mentation. It was during this interval that the present topographic

detail in its large phases as now expressed on the West Slope was deline-

ated.

From the data at hand, the successive steps seem to have been as fol-

lows: At the close of Cretaceous sedimentation and voleanic activity the

West Range of the Andes was elevated; the west side of the uplifted sec-

tion was subsequently worn down to a poorly graded surface, at least in

the Huacho-Oyon section; this stage was followed by a strong uplift and

subsequent deep dissection, or erosion. This was followed, in turn, by

the depression of at least the shoreward edge of the post-Cretaceous land-

surface and the initiation of Tertiary deposition. In other words, Ter-

tiary deposition took place on the partly drowned edge of the West Slope.

Ocona-Cora Puno Section.—This section has been chosen for the rea-

son that the observed facts show considerable variation from the one

already described. The town of Ocofia is situated on the coast some

seventy miles to the northwest of Mollendo. The point known as Cora

Puno is located near the western edge of the West Range. It is one of

the highest collection of voleanic peaks on the west of the Cordillera.

Starting at Ocona, we shall run our section across the immense plain or

desert of Cuno cuno to the valley of the Chorunga, thence up the Andaray

cuesta to Cora Puno and the West Range.

The formational and topographical facts are roughly shown in the

accompanying section. At Ocofa, we have the pre-Tertiary coastal ridge

facing the present shore line. On its west slope may be seen patches

of Tertiary and post-Tertiary still inviting the attack of the Pacific

waves. Passing inland, however, we find that, as we reach the level of

the pampas, pre-Tertiary formations may be seen at many places sticking

through the thin sheet of late sediments at elevations of 3,000 or more

feet above sea level. As we pass towards the north we soon realize the

fact that the surface of the Cuno cuno plain has once been covered in part

at least by a sheet of mud-like lava. Only in the deeper: side-valleys

leading to the Ocoha canyon can we see the sediments as recognized near

the coast. This light-colored sheet now persists until we reach the steep

slope that passes into the Chorunga Valley. Here the Chorunga as well

ANNALS N. Y. ACAD. SCIENCES VOLUME XNII, PLATE XXXI

Iie. 1

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PLATE XXXI

INNER EDGE OF THE PAMPA CUNO CUNO

Ite. 1.—A view of the inner edge (inface) of the Pampa Cuno cuno (coastal

plain) as seen from the stripped upland of the west slope in the region of the

Valley of the Choringa-Ocona-Cora Puno Section.

CUESTA DE IQUIPI, NEAR VIEW

Fre. 2.—A closer view of Cuesta de Iquipi, or inface of the coastal plain as

seen from opposite the Rio Grande Valley-Ocona-Cora Puno Section.

Corro Puno

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES QA

suse eo ete

[<<*] Diorite

Cretaceous

RSs Sandstone and

[&2%e"] Lavas and Volcanic Ash of Corro Puno

BSS] Basalt

*Syay] Porphyry

Ti Mud Lava

Tertiary

=| Sands

Shales

Wie. 1.—G@ecological section from Ocona to Corro Puno

as the Rio Grande and the Ocona have succeeded

in cutting through, not only the soft lava referred

to above and the underlying sediments, but also

have deeply incised themselves in the underlying

pre-Tertiary formations of the partly buried foot-

hills, or, more accurately stated, have cut through

the coastal plain formations until in the Valley

of Chorunga the line of erosion adjusted itself to

a pre-Tertiary topography. That is to say, the

Chorunga simply cleaned out the mass of sedi-

ments and sheet of lava which had been dumped

into it during Tertiary time. This is evidenced

by the patches of lava that still hang in the little

recesses on the southwest slope at various points:

between the Rio Grande and the Andaray cuesta.

Ascending the Andaray cuesta, we go but a.

short distance up the slope before we again see:

remnants of the familiar lava sheet of the Cuno

cuno plain, still nestling on the slopes and in the

protected valleys of the uncovered foothills lead-

ing up to the base of the Cora Puno domes. At

Andaray and Yanaquiqua, we can see the same

facts well illustrated. Andaray is located in a

depression in the foothills, the bottom of which

is still in part occupied by the light-colored lava.

From Andaray, we pass up another steep slope,

or series of slopes, composed in part of pre-Ter-

tiary rocks, only to find that we are again on a

somewhat dissected plain which extends away to

the east in the direction of Chuquibamba and

north in the direction of the picturesque Cora

Puno. Lithologically, this is the same type of

rock as that which we have already seen in the

Chorunga Valley, the Andaray cuesta and the

inner edge of the coastal plain. It is my belief

that the plains at the base of Cora Puno group

of domes are but parts of the same lava plain that

we have already recognized in a portion of the

Cuno cuno desert. Continuing our journey to

the north, we come to the somewhat rugged,

jagged edge of a still higher plain that is made

949 ANNALS NEW YORK ACADEMY OF SCLENCES

up of a vesicular scoriaceous black to gray lava. Standing on top of this

is the picturesque group of Cora Puno domes, four in number. So far

as personal examination was made, it was found that lower slopes of the

domes were composed of a succession of ropy lava flows alternating with

voleanic ash and cinder-like layers. Nestling in the valleys between the

domes are fine glaciers. On the southwest side of the group, there may

be seen a very fine icefall, with an exposed edge of not less than 200 feet.

At the time of visiting this locality, the domes were well covered with

snow. So symmetrical and smooth did the domes appear, even to their

summits, that the writer felt that with a specially constructed snowshoe

an ascent might be made without serious difficulty.

The base of Cora Puno is about 14,000 feet above sea level. The domes

are not less than 20,000, and one of them may reach the 21,000-foot

mark.

Before interpreting the physiographic value of this section, let us’ look

at another running from Ocofa to Caraveli. In this case we start at

the coast with the nearly buried coastal ridge facing the Pacific. Going

inland we pass over a succession of plains arranged in bench-like order.

The natives have applied names to each of these. Near the coastal ridge,

the plain is composed of sedimentary formations, but we do not go far to

the interior before we recognize the fact that the light-colored lavas form

the only visible lithological unit exposed in the deeper valleys of the

plain. Should we go from Caraveli to Atico, we will again cross the lava

plains, but in the lower part of the Atico group of valleys we will see that

sands and conglomerates become important members of the coastal plain

deposits. Furthermore, after we leave the outliers at the shore line of

Ocona, we do not come again in contact with the pre-Tertiary oldland

until we pass through the Quebrada de Chin Chin to the Caraveli Valley.

From Caraveli inland, the foothills ascend rapidly to the elevation of the

basal plain of Cora Puno. Above this elevation tower, here and there,

occasional peaks, most of which are quite different in topography from

those of Cora Puno. They represent elaborate groupings of spires and

pinnacles, at places so steep that snow seems unable to cling to their sides.

Should any one of the enthusiastic mountain climbers who have recently

achieved noted success in mountain climbing in central Peru care to in-

vestigate this region, he will be sure to find opportunities for testing his

skill, of which he had never dreamed.

We see, then, in the first-mentioned case, the oldland, or a portion

of the West Slope, has scarcely been buried beneath the Tertiary deposits

and no small part of these is composed of mud flows capping the entire

series. In the section to the west, however, we have seen that sedimenta-

DOMES OF CORA PUNO

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ANNALS N. Y. ACAD. SCIENCES VOLUME XNII, PLare XXNII

Fie. 1

Fig. 2

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 943

tion was predominant near the edge of the coastal chain and also in the

lower part of the Atico Valley; but it should be added, in the latter case,

that much of the upper part of the sediments may be post-Tertiary and

associated with glacial waste. At least, in the Atico-Caraveli section, we

see nothing of the oldland upon which these late sediments rest, until we

approach the vicinity of Caraveli. From these sections, it would appear

that the West Slope in this locality was never reduced, or worn down to a

grade, whereby even an approach to an even sky-line was attained, as

noted in our northern section, but was occupied by valleys and hills of

. large dimensions when Tertiary sedimentation and volcanic eruption be-

came prominent phenomena. The pre-Tertiary foothills were only partly

buried in sediments, but later they became almost obliterated by the ac-

cumulation of mud flows of great thickness and areal extent.

It is thus evident that the physiographic history of the West Slope is

by no means simple. To get a clear insight into the possible variations

and succession of events, sections should be studied in every department

and province facing the Pacific coast.

In the main, however, I believe that we have noted the chief succession

of events and the principal factors involved, namely:

(1) The development of an oldland surface upon Cretaceous and

probably older formations and corresponding to the western slope of the

West Cordillera.

(2) Its elevation and deep incision of valleys.

(3) The depression of the shoreward portion, which, during Tertiary

time, was in part at least covered with Tertiary deposits, and portions

of which now form the present coastal plains.

(4) The differential elevation of the coastal plain deposits and the

extension of the drainage of the oldland to the present coast line. In

all probability, no small part of the original area of Tertiary deposition

may now be beneath sea level.

HIGHLAND PLATEAU OR INTERCORDILLERAN BELT

To the belt of country lying between the eastern and_western ranges of

the Andes, I have applied the terms Highland Plateau or Intercordilleran

Belt. These terms are used in the broader sense and for the need of

better ones. Although presenting a complex geologic history, it can be

traced from central and northern Bolivia throughout the entire length of

the Peruvian Republic and into Equador; its width, in general, varying

from 100 to 200 kilometers.

DA 4. ANNALS NEW YORK ACADEMY OF SCIENCES

In the south of Peru, it is in part occupied by the most picturesque

inland body of water of which the South American continent can boast,

the well-known Lake Titicaca. To the southwest of Lake Titicaca, we

have a somewhat dissected area drained by the Rivers Blanco, Canco-

marca and Maure. To the southeast, we find the extension of the broad

plains leading to La Paz, Bolivia. To the northeast, the Titicaca plains

soon melt away in the foothills of the East Cordillera, whose long un-

broken front, as seen from Lake Titicaca, presents a spectacle of moun-

tain scenery that is not duplicated until we reach central Peru. To the

northwest, we find that the Titicaca plains give place quite rapidly to a

highly dissected belt. The streams coming to the Titicaca basin from the

northwest are short, and their floors are very much ageraded, representing

arm-like extensions of the enormous pampas to the north of the lake. As

soon as we pass over the drainage divide at La Raya, we drop into an

area that has been deeply cut by the drainage system of the Apurimac.

It represents the headwaters of the Tambo and Ucayali, the latter join-

ing the Marafion, and it, in turn, emptying into the Amazon. Of this

stretch of country, the writer is personally acquainted with the Cuzco

section, or rather, the belt etxending from La Paz, Bolivia, to a cross

section through Cuzco.

Let us search for a locality where we can get a clear sweeping view,

both to the northeast and southwest. To do this, we must ascend to the

upland surface into which the Urubamba and Apurimac systems of drain-

age have deeply incised themselves. Various points of vantage may be

found in the highlands overlooking the valley of the Urubamba be-

tween Cuzco and the Pueblo Urubamba, or from high points overlooking

the valley of the Apurimac. Selecting a high point among the latter, and

looking to the northeast, we will see, in the foreground, as it were, reall

two ranges looming up before us, and far above the level on which we are

standing, one lying between the Apurimac and the Urubamba, and the

other between the Urubamba and the head-waters of the Madre de Dios.

While topographically these form parts of the East Cordillera, I am

inclined to believe, from a geological point of view, that only the range

lying northeast of the Urubamba-Villcanota valleys belongs in the Hast

Cordillera proper. One could not wish for a more varied bit of mountain

scenery than that from Sicuani to Cuzco. To the northeast, we have the

pinnacle, spire and-knob-like topography that characterizes a region of

great volcanic activity. To the southwest, however, we have a very dif-

ferent expression, that which characterizes a belt occupied largely by

sedimentary formations, here and there disturbed by local volcanic in-

trusions.

PLATE XXXIII

INTERCORDILLERA NEAR CUZCO, LOOKING EAST

Fie. 1.—A view of the sky-line of the intercordillera belt in the region of

Cuzco, looking east.

INTERCORDILLERA NEAR CUZCO, LOOKING SOUTH

Fig. 2.—A view of the sky-line of the intercordillera belt in the region of

Cuzco, looking south.

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ANNALS N. Y. ACAD... SCIENCES VOLUME XNII, Phare NXNIII

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MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES Q45

From the point of view selected, we see a billowy mass of ridges whose

summits correspond roughly to the level of those on which we are stand-

ing. This view passes across the Intercordilleran Belt, while the former,

in part, belongs to the East Cordillera. From a physiographic point of

view, we thus have, first, the Intercordilleran Belt, a highly dissected

plateau, longitudinally drained by the Apurimac and its branches head-

ing against the east flank of the West Cordillera; second, the East Cor-

dillera, behind which run the Urubamba and Villcanota systems, also in

longitudinal position ; but these finally succeed in cutting their way across

the final range to join the Ucayali, which passes west of north through

the Department of Loreto and drains for many miles the inner edge of

the Amazonian plains. It is on this river that the important inland city

of Iquitos is situated.

Cerro de Pasco Section —Let us now examine the topographic charac-

teristics to be observed in a cross section passing through the mining town

of Cerro de Pasco and connecting with that which we have already de-

scribed under the name of the Huacho-Oyon section. In this way, we

can get a long-distance view of the principal features of these geographic

provinces extending from the coast to the East Cordillera. As we have

already stated, Oyon stands on the western slope of the West Range. Let

us now go from Oyon to an advantageous point of ‘view in the Uchuc-

chacua section, where we may obtain a clear view both to the east and to

the west. Such a point may be found near the main trail from Oyon to

Cerro de Pasco and not far from the “pass,” with an elevation between

16,000 and 17,000 feet above sea level. To the west and southwest will

be seen the western slope of the West Cordillera, as already described.

To the east, we will see, some 3,000 or 4,000 feet below us, a stretch of

rolling undulating country extending to and beyond Cerro de Pasco.

Occasional knobs and ridges may be seen at various points standing on

the plain and serving as guides and mileposts to the explorer. From the

same point of view, on a clear day, the East Range may be seen, away

to the east and northeast of Cerro de Pasco. Here may be found the

grandest and most magnificent group of ridges and peaks rearing their

summits far above the line of permanent snow. There are peaks here

that pass far above the 20,000-foot level. Into this Intercordilleran

Belt the headwaters of the Huallaga have cut their way southward into

the vicinity of Cerro de Pasco, having pushed through a part at least of

the Hast Range. It is important to note here that the Huallaga did not

succeed in cutting its way through the entire Hast Cordillera to join the

Ucayali, but was forced to maintain a longitudinal position as far as

Huanco. Then it broke through the East Cordillera to join the Marafion

IAG ANNALS NEW YORK ACADEMY OF SCIENCES

far to the north. It thus maintains a position approximately parallel to

the Ucayali throughout a long distance.

The Cerro de Pasco plain is quite clear as far south as Oroyo. In this

part of the plateau we again find a pretty sheet of water, but very much

smaller than the inland Titicaca. From this body of water, known as

Junin, the drainage, instead of going to the Huallaga, now goes to the

southeast, the main line of drainage being the so-called Mantaro. From

Oroya to Huancayo, the Mantaro has made a most picturesque valley, on

the floor of which are located many prosperous agricultural communities.

Ascending the highland in the region of Huancayo, we can see to the east

the picturesque Cordillera de Marca Valley, really a part of the Hast

Cordillera, while to the west may be noted the West Range presenting an

almost unbroken front.

From Huancayo, the Mantaro continues its course to the region of

Mayoc, where it breaks through the frontal East Range and deflects again

to the northwest for some distance, but finally elbows its way across the

last eastern barrier near Huaribamba and joins the Apurimac. We thus

find here another illustration of a pattern of drainage which suggests an

adjustment to structure, such as may be seen in many other regions the

world over.

Relative to the section between Mayoc, or the first elbow of the Man-

taro and the Cuzco-Abancay region, I may add, from data obtained from

explorers, that it 1s apparent that it is a broad highland belt into which

the east tributaries of the Apurimac have incised themselves until they

have cut the entire section into a veritable labyrinth of canyons. In

other words, it is simply a continuation of what we noted to the southeast

of Cuzco.

Casma-Huaraz-Huacaybamba Section —Let us now look at a section

through the northern part of the Department of Ancachs beginning

with Casma on the coast and ending at Llata on the Marafion. At

Casma, and far to the north and south, the foothills of the west slope

form the coast line. The main transverse valleys have been aggraded near

the present shore line, but we do not go far to the interior before the flat

floors are replaced by steep-sided V-shaped valleys. Here again we have

the rapidly descending surface of the West Slope; to the interior, this

culminates in the West Range, which in turn overlooks the valley of the

Huaraz. The highest points of the range in this section do not exceed

15,000 feet. From an examination of the Raimondi map, it will be seen

that it is well defined throughout a large part of the Department of An-

cachs. Note the fact, however, that in the place of the broad plain, as

found in the Cerro de Pasco section, occupying the belt between the Hast

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 247

and the West Cordillera, we have a comparatively narrow valley running

parallel with the West Range. It is drained by the River Huaraz, occu-

pying a longitudinal position throughout some three provinces, and finally

in the Province of Pallasca turning to the west to join the Santa, which

empties into the. Pacific.

Continuing our section to the east from Huaraz we find in the place of

the deeply dissected intercordilleran plain another range of enormous

proportions and running parallel with the one already described. It

teaches elevations of more than 20,000 feet at various points. From its

east slope, however, we now see the much-dissected Intercordilleran Belt

extending to the valley of the Marafion, a distance of perhaps seventy

miles. We thus have in this section two ranges on the west side of the

Intercordilleran Belt, instead of one, as in other sections noted; further,

instead of an intercordilleran lowland, as noted in the Cerro de Pasco sec-

tion, we have a deeply dissected belt whose uppermost surface has a

marked descent towards the valley of the Maranon. To the east of the

Maranon, we again have a somewhat softened expression of a continuation

of the East Cordillera. :

Let us now consider the position of the Maranon drainage system. It

heads to the south in the Department of Junin in a collection of lakes

within the limits of the Intercordilleran Belt. Following the main line

of drainage, we find that it, like many others, skirts the east edge of the

intercordilleran highland for many miles to the north, and finally, in the

Department of Amazonas, turns east to join the Amazon system, of which

it is an important member. Note again the longitudinal position of the

Marafion with reference to the geographical provinces as outlined.

To the east of the valley of the Maranon, the Raimondi maps show a

fairly well developed range separating the drainage of the Huallaga from

its western neighbor. Just what this range is, geologically, is unknown

to the writer, but it is believed to correspond to the East Cordillera.

Piura-Huangabamba Section——To the east of the Piura-Salitral River

the long western slope is absent, and we encounter immediately the con-

tinuation of the West Range with a rather abrupt slope, though softened

in its topographic expression, as compared with sections to the south.

This is followed by the intercordilleran highland, already thoroughly cut

to pieces by parts of two systems of drainage. The section of the Inter-

cordilleran Belt between Huangabamba on the north and Chota on the

south is drained by the rivers Huangabamba and Chotana. These be-

come confluent in the river Chamaya, which empties into the Marafion.

Note again the longitudinal position of the Huangabamba and Chotana

rivers with reference to our geographical provinces, while the Chamaya is

typically transverse in position.

D48 ANNALS NEW YORK ACADEMY OF SCIENCES

In the north part of the Department of Piura, we have some unex-

pected topographic variations. By consulting the Raimondi map, it will

be seen that the part corresponding to the West Range has projected a

long spur to the southwest, known as Cerro da La Brea, or Amotape

Mountain. It is composed of Cretaceous shales and dioritic intrusives.

It is flanked on both sides by the coastal plain formations. Note also

the arrangement of drainage between the spur and the headwaters of the

Huangabamba. The rivers Chira and Catamayo represent the main line

of transverse drainage, while its tributaries the Suipira, Quiros and

Macara represent the longitudinal drainage of the somewhat depressed

section of the intercordilleran plain.

HAST RANGE AND EAST SLOPE

Concerning the East Cordillera, we have already seen in our various

views that it forms a prominent range of enormous altitude at various

points and maintains its identity throughout the entire length of the

Peruvian Republic. Its highest altitudes exceed those of the West Range.

The most picturesque views may be had from the region of Lake Titicaca,

where, on a clear day, it may be seen passing north from Bolivian terri-

tory into Peru and presenting apparently an unbroken front, as far as

the eye can reach, to the northeast. It maintains its identity throughout

the departments of Puno and Cuzco, reaching enormous altitudes in the

Department of Junin, where it is deeply cut into serrated forms. We

then can follow it through the department of Huanuco and San Martin

and finally into the Department of Amazonas, where it apparently sep-

arates into two ranges and continues as such into Ecuador. While no

great elevations are attained in northern Peru, they assume again in

central Ecuador their old-time grandeur.

Concerning the East Slope of the East Range, I regret very much that

I had little opportunity to see enough of this geographic province to war-

rant personal description in any detail. Only in southern Peru have I

penetrated the East Cordillera to a point where a far-reaching view of

the foothills and, beyond these, the great stretch of rolling and undulat-

ing lowland, may be had. The foothills proper do not occupy a very wide

belt. They quickly descend to an elevation of not more than 4,000 feet

and probably less, where, from the long-distance view at least, one would

Judge we should encounter the inland edge of the great Amazonian plains.

These occupy the east portion of the Department of Puno, a very large

part of the Department of Cuzco and nine-tenths of the Department of

Loreto. It is on these eastern slopes and the huge plains below that we

find uncivilized tribes of Indians, or the native “salvaje.”

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 24.9

TopoGRAPHIC EXPRESSION AS RELATED TO THE GEOLOGY OF THE

PERUVIAN ANDES

PLAINS AND SECTIONS

Zorritos-Lambeyaque Plain.—The formations of this division of the

coastal plain are, so far as known, Tertiary in age. They are entirely

sedimentary and are composed in the main of clays and sands with occa-

sional pebble and conglomerate beds. From a section prepared by me.

between Fernandez and the shore line, something like 3,000 feet of sedi-

ments can be calculated. More than 2,000 feet have been penetrated by

the drill in search of new petroleum-bearing horizons both in Zorritos

and Lobitos. I wish here to emphasize the point that the only localities

where a topographic expression resulting from uplift and subsequent

erosion may be found are confined to the three localities, Zorritos, Lobi-

tos and Nigritos, where petroleum in large quantities has been obtained.

Each one of these places is located on the east limb of a somewhat broken

or locally faulted anticline. ‘The formations as seen in the Fernandez

section are regarded as lower and middle Tertiary. In the Lobitos, it is

believed that we have middle and upper Tertiary.

In the Paita section, reference has already been made to the occurrence

of a series of sands and conglomerates resting unconformably upon a

mass of red clays. These deposits contain some fossils, apparently the

same as those living at the shore line to-day. They are now 250 to 300

feet above sea level. As we pass in the direction of Piura, we can see

little mesa-like elevations. ‘These were found to be composed of the red

clays seen in the lower part of the Piata section. It is thus evident that

the red-clay deposits were elevated and eroded before the deposition of

the conglomerates referred to above; and, further, the erosion must have

been largely confined to the outer half of the Paita section.

It is important to note another fact at this point. At a number of

places in the Paita-Piura Pampa or Despoblado, as the natives are accus-

tomed to call these plains, you cannot fail to see small areas strewn with

shells, all of which appear to be specifically the same as those living on

the present shore line. This would suggest that, in very late geological

time, the Paita-Piura Plain was beneath sea level, and that it was subse-

quently elevated to its present position.

As we pass to the south, the despoblado has been barely scratched by

the Piura River and the surface subsequently littered up with wind-blown

sands. That is to say, topographically, the plain is so young that it yet

has the same expression as when it emerged from sea level. This con-

250 ANNALS NEW YORK ACADEMY OF SCIENCES

dition is maintained throughout the Desert of Sechura and the Despo-

blado de Olmos. From Chiclayo and Etén south, we have but a rim of

the coastal plain left. It attains considerable proportions in the mouths

of the valleys of Pacasmayo, Chicama and Santa Catalina. From a topo-

graphical point of view, we have here but a narrow expressionless plain,

with its seaward edge rapidly retreating under the attack of the Pacific

waves.

The two prominent points where topographic expression relieves the

monotony of the plains are Cerro or Silla de Paita and Cerro de Yllesca.

These are, in a word, half buried outhers of the foothills of the West

Cordillera. Similar cerritas may be seen in the broad opening of the

Pacasmayo Valley. All these were little islands in a Tertiary sea.

They are composed of Cretaceous shales and sandstones which have under-

gone metamorphism under the effect of intrusives.

Chincha-Olmos Plain.—From Chincha to Pisco, we have the same

monotonous plain. It is not until we reach Pisco that we find a little

topographic relief and this time again associated with the uplift of a

series of light gray to cream-colored clays. We need, however, only to go

a little distance inland to see a continuation of the pampa in the direc-

tion of Ica. Should we follow the coast, we should find that a consider-

able area is occupied by the outliers of the foothills, but these are quite

modest in topographic expression. These pre-Tertiary hills continue to

the mouth of the river Ica and attain considerable width. ;

I wish to turn aside here for a moment to refer to the Peninsula of

Paracas, a short distance south of Pisco, since its geology is somewhat

unique. Some time ago, coal was found in the cliffs of the peninsula,

and a company was formed to exploit the deposits. Examination of

the waste brought out of some of the prospects revealed the occurrence of

true Carboniferous plants. It is the only locality known to the writer

where undoubed Carboniferous and coal-bearing measures occur. So far

as my observation goes, the Peninsula of Paracas is by far the most

ancient “morro” on the entire coast of Peru. It is my belief that the

formations entering into the remainder of the coastal chain, or Cadena

de la Costa, are, geologically, much younger.

Passing to the other side of the shore ridge of which Paracas is the

northwestern extension, we find the lowland facing the actual foothil!s

of the West Cordillera cut into by the Ica River. Here we are relieved

to find another good section of the same light-colored clays as seen in the

Pisco section. As soon as we reach the Pampa de Huayuri, we are again

greeted by an enormous stretch of high plain, and its monotony is only

relieved when we reach the modest canyon cut in it by the Rio Grande.

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES O51

On the southeast side and close to the coast is the Cerro de Yungi, an-

other pre-Tertiary outlier. From this point to the termination of the

Chincha-Olmos section we see again just the young Tertiary plain wigs y

eut into by the rivers de Acari and Yuaca.

Ocona-Moquequa Section—In the southern section, which may be

aptly termed the Ocona-Moquequa section, we have a repetition of the

same topographic expression and the same formations involved, with one

additional feature, and that is the role played by the mud flows. As

already indicated, they form avery large part of the formations entering

into the coastal plains extending from the Valley of Caraveli to the

Pampa de Clemisi. The special feature of the south section is the per-

sistence and strong relief of the coastal chain. It extends practically

throughout the entire length of the coast line. I have indicated that

the formations entering into the structure of the Cadena de la Costa are

the same as appear in the north. While this may be true in some paris,

I am inclined to think that, in the south, other formations than those

of the foothills of the cordillera so far noted may be found in the Cadena.

de la Costa.

Huacho-Cerro de Pasco Section.—t\ have already called attention to

the occurrence of an evident west slope facing the Pacific Ocezn and the

coastal plains. It has a marked inclination towards the sea. It is thor-

oughly cut to pieces by a network of transverse valleys. It is only when

you ascend the uppermost edges of any of these valleys that this feature

becomes apparent. Nevertheless, above the general sky line of the slope,

we can see many elevations.

The geology of the above section is as follows. Near the coast the

formations are sandstones and shales into which have been intruded two:

types of volcanics. These are then followed by a broad band of crystal-

lines, probably diorites and related types. These are, in turn, followed.

by shales and sandstones. Large coal deposits are to be found in this

part of the section. The shales and sandstones extend to and beyond

Oyon. In the vicinity of Ututo, the sandstones and shales are replaced

by an enormous thickness of limestones. Into the limestones and the:

inner edge of the sandstone and shale formations, enormous volcanic

masses of at least two kinds have been thrust. In this section at least,.

the volcanics form most of the crests of the West Cordillera. The sedi-.

mentaries, especially in close proximity with the volcanics, have been

folded and crushed on a very large scale.

The eastern slope of the West Cordillera is remarkably well defined.

Limestones are here turned up on edge, and adjacent to this horizon we-

find the volcanics. Each formation presents its own type of topography.

952 ANNALS NEW YORK ACADEMY OF SCIENCES

At some points along the eastern slope, the escarpment is of such a char-

acter as to suggest faulting on a very large scale.

Continuing across the Intercordilleran Belt, consisting of an undu-

lating plain, some 3,000 to 4,000 feet below the summits of the West

Cordillera, we find the formations involved in its structure to be sand-

stones, shales and limestones, through which, at various points, knobs of

voleanic rock have pushed their way and now form a part, at least, of the

principal relief of the Cerro de Pasco lowland. The town of Cerro de

Pasco is built on the slope of one of these knobs. Here we find a mass

of voleanic rock in contact with a large body of limestone. It is on or

near the contact that the famous ore body is located. Going northward

over the intercordilleran lowland to Goyllarisquiseca we again encounter

coal-bearing formations. Just what are the stratigraphical relations be-

tween these andthe coal-bearing measures of Ututo and Cajatambo is

not known. Beyond this point the writer has not penetrated the wilds

of the East Cordillera. From data obtained from prospectors and en-

gineers, I have reason to believe that the principal formations involved in

its structure are very much older than any we have seen in the West

Cordillera. Probably Devonian and Silurian and older terranes asso-

ciated with a huge mass of intrusives make up the great part of the Hast

Range. By way of comparison, I may add that the formations found in

the Lima-Oroya section duplicate, in the main, the Huacho section, both

in succession and kind. Oroya stands at the south end of the Cerro de

Pasco lowland. Here we have the expected limestones and shales and

associated intrusives noted to the north.

Ocona-Cora Puno Section.—Starting at Ocofia, we meet first of all the

outhers within the limits of the coastal plain, or the Cadena de la Costa.

Ascending the shoreward escarpment, we pass over the ridge and on to

the edge of the enormous pampa which we already know under the name

of Cuno cuno and have recognized as an intergral part of the Coastal

Plain province. Should we pass into the canyon of the Ocona, we should

see the Tertiary sediments resting unconformably upon a series of sand-

stones and shales. Inland along the line of our proposed section, the

Tertiary sediments give way to the development of an enormous mud

flow, which, on the inner edge, or escarpment overlooking the valleys of

the Chorunga and Ocofia, is not far from 1,000 feet in thickness. In

the canyon of the Ocofa, in the region of Piuca, the pre-Tertiary sedi-

ments recognized nearer the coast have been largely replaced by various

types of volcanics. There are at least three types to be found in the

Oconha and Chorunga valleys, namely, a dioritic, a trachytic and a basaltic

type. Their succession of volcanic activity was probably in the order

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 953

named. From the valley of Chorunga to Andaray, the dioritic forma-

tions constitute the principal formation. Ascending the highland above

Andaray, we again find a broad plain extending away to the southeast in

the direction of Chuquibamba. The plain is composed of the same sort

of mud flow as we have seen on the inner escarpment of the Coastal

Plain overlooking the Chorunga Valley. Through this protrude various

knobs of diorite, the same horizon as we saw in the trail from Chorunga

to Andaray. Into this plain the west tributary of the Majas River chas

cut a deep valley not only through the mud flow, but well into the under-

lying crystallines. On this floor rests the city of Chuquibamba.

Continuing our course to the north and northeast, we come finally to

the edge of the lava flows forming a veritable platform on which were

built the four confluent domes of the Cora Puno composed of lava, ash

and scoria. A very large part of their slopes is above permanent snow

lne. These domes undoubtedly surpass 20,000 feet in altitude. From

the northeast slope of Cora Puno we have before us a gently ascending

plain, with cerros appearing here and there above the general sky line.

Among these are the majestic Solamana, Leon Wachang and others with

unpronounceable Indian names. And here let me say, if any enthusiastic

mountain climber wishes to test his real ability, he should not miss trying

the spires of Solamana. The plains between the peaks are, in part, at

least, made up of a light-colored mud flow, lithologically the same as that

seen on the other side of Cora Puno and the inner edge of the Coastal

Plain or the Iquipi cuesta. Occasional small knobs of limestone protrude

through this sheet. Also, where some of the streams have cut to any

great depth, we sometimes find limestone exposures. That is to say, the

light-colored lavas, the dark basal lava platform and the superimposed

cones of Cora Puno, probably rest upon a floor of limestone on the north-

east and dioritic crystallines on the southwest of Cora Puno. Such of

the smaller spire-like hills as were examined were found to be volcanic.

In a word, then, comparing our sections, we shall find that we have a

similar succession and order, with the exception that in the Huacho-Oyon

case the Coastal Plain is absent. Physiographically, Cora Puno and the

series of snow-covered domes to the southeast and northwest are situated

well up the West Slope and not far from the West Cordillera. The im-

portant mining camp, Caylloma, is located on the east slope of the West

Cordillera, into which the headwaters of the Apurimac are now cutting

their way.

To the northeast from Caylloma to the Cuzco region we have the broad-

ened Intercordilleran Belt, literally cut to pieces by the labyrinth of val-

leys oceupied by the various tributaries of Apurimac and the Villcanote.

O54. ANNALS NEW YORK ACADEMY OF SCIENCES

To the southwest from Cora Puno, we have the West Slope extending

to the edge of the Coastal Plain, then follows the Coastal Plain to the

Cadena de la Costa, the latter now bordering the present shore line. —

If time would permit, a section passing from Mollendo through Are-

quipa to Puno could be shown to duplicate,.in the larger phenomena, the

facts already brought out in the Ocofa-Cora Puno section. I wish, how-

ever, to say just a word concerning the Titicaca region. he lake occu-

pies a portion only of the Intercordilleran Belt. Within this basin, Ter-

tiary sediments have been deposited. There is reason to believe that the

Titicaca basin represents an area the depression of which was associated

with down-faulting on a large scale. It probably extended from the

north end of the Tertiary Titicaca well down to La Paz. Further, the

Tertiary deposits rest upon limestones and shales. We are probably war-

ranted in correlating the latter with the limestones and shales of the

Cerro de Pasco lowland. To the west of the lake, we have the shales and

limestones extending to the divide of Cerros de Toledo, where we again

come in contact with voleanic intrusives, the true core of the West Cor-

dillera. The belt of limestones and shales on the east slope of the West

Cordillera has been traced to the Cuzco section. It physiographically

belongs to the Intercordilleran Belt.

The section of Forbes brings to light the same physiographic features,

the Coastal Plain, the West Slope and West Cordillera, followed by the

Intercordilleran Belt and finally the East Cordillera.

NoTEs oN Harty MINING IN PERU

To attempt to discuss the mineral resources of Peru in detail is not my

intention at this time. I wish, however, to say one word relative to the

early history of mining, its initiation by the Incas, its subsequent devel-

opment during the period of Spanish rule, and finally to present a brief

geographical and geological correlation and distribution throughout the

West Cordillera and the Intercordilleran Belt.

While the Incas as a race were decidedly agricultural and pastoral in

their vocations, they were, nevertheless, not ignorant of the use of the

precious metals. This is proved by the occurrence of gold and silver

vessels discovered in their notable monuments, known under the name of

“huacos.” The huacos are large quadrangular and pyramidal earth-

works. They were probably used in connection with religious rites and

ceremonies. While these constructions, or monuments, may be counted

by the score near the coast, and usually are located on the floor of the

broader valleys near a locality affording protection, the best preserved

MARSTBERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 955

examples seen by the writer are to be found in the Valley of Santa Cata-

lina (Salaverry). Very fine huacos may also be seen near Lima and

within a day’s ride from the Capital. Quite recently a pair of vases,

each apparently beaten out of a solid piece of gold, was found in a

hhuaco near Lima. From the Valley of Nasca, I have seen a large col-

lection of gold bands, undoubtedly used as wristlets and as ornaments

for the head. All these were discovered in the’ interior of a huaco. Hence

all must have been of Inca manufacture, and from the crude metal ob-

tained from its original source by these people.

With the invasion of Pizarro and his followers, and the subsequent

establishment of Spanish rule over the Inca people, we have to note the

introduction of a new regime. The invaders were primarily interested in

the discovery and accumulation of the precious metals. On the other

hand, the native was agricultural and pastoral in habit. As soon as the

Spaniard had gained a knowledge of the gold-bearing possibilities of

their newly acquired territory the labor problem became an important

one. There could be but one outcome or solution on the part of the in-

vaders. The natives were thus pressed into mining services and driven

away from their chosen vocations. For a period of something like three

centuries the native remained in a state of servitude to the Spanish rulers

and people. If we can put any credence in various sources of informa-

tion concerning the suffering of this inoffensive race, it seems most re-

markable that they did not revolt and make at least one heroic effort to

free themselves from the servitude into which they had fallen. It was

‘during the period of the Viceroys that the gold-bearing resources of Peru

became legend. That Peru was rich in the yellow metal was evident to

the invader Pizarro upon his first survey of Cajamarca. Here he dis-

covered and took charge of gold and silver to the amount of some three

millions of soles in actual value.

To speak in general terms of the mining activities under Spanish rule,

we may conveniently group the localities of maximum activity in the fol-

lowing manner:

(1) Cajamarca-Pataz Section. In this region, old Spanish prospects

may be counted by the score in numerous valleys.

(2) Huaraz-Cajatambo-Cerro de Pasco Section. While the old Spanish

workings are not yet known to be as numerous in this section, it is never-

theless certain that large amounts of both gold and silver were obtained,

especially in the region of Cerro de Pasco.

(3) Cotahuasi-Andaray Section. In this section and as far west as the.

valley of the Chala, there are many abandoned prospects. In Andaray

and the Cotahuasi vicinities, as well as in the Cerro de Pasco region,

much work has been done since the establishment of the republic.

256 ANNALS NEW YORK ACADEMY OF SCIENCES

(4) Cuzco-Cotabamba Region. The region of Cotacamba and adjacent

valleys was the scene of great mining activity in the early Spanish days.

Many of these prospects have likewise been worked in late times.

(5) Poti-Sandia Section (East Cordillera). This is known to con-

tain not only a large area of mineralized territory—gold-bearing quartz

veins—but also an abundance of placer on the eastern slope of the Hast

Cordillera. :

(6) Huanaco Section (Hast Cordillera).

Just a word as to the distribution of the principal mining localities

and their relation to the geology of the Cordillera. After seeing a large

number of ore-bearing sections in the south, center and north of Peru,

the relation and association of zones of maximum mineralization with

certain formations becomes very clear indeed. Let us return for a moment

to our Huacho-Cerro de Pasco section. Near the coast, there are a few

intrusives which have pushed their way through the sandstones and,

shales. Apparently associated with these volcanics are gold-bearing

veims which have been prospected from time to time. Nothing of very

great value, however has been found here. The Spanish prospector did

not find this little crop of veins very attractive. He was not slow to hunt

more pleasing ground further to the interior.

It is not until we pass to the zone of the West Cordillera, where there

are enormous intrusive bodies bordered by limestones and shales, that we

find ore-bodies of large dimensions. On the west side of the Cordillera,

we have a group of silver-copper-gold veins, some of which can he

traced for more than a kilometer, with widths approaching 20 meters.

Should we pass over the divide to the East Slope, from which we see the

Cerro de Pasco lowland, we will find both on the slope and in the valleys

leading to the crests of the Cordillera another group of veins that are

undoubtedly associated with the east contact of the intrusives with the

limestones and shales.

Throughout this region may be seen many old plants (arrastres) and

the still more ancient quimbolete, where the ores were treated for the

recovery of gold and silver. The amount of visible tailings show to what

extent the early prospectors worked. Since the foundation of the repub-

lic, the native has likewise continued to work in this region, but, of course,

in the old-fashioned way. It is not at all uncommon to find Indians

in possession of solid silver utensils hammered out of a single piece of

silver. Should you visit the plazas of any of the villages in these sections,

you would find the silversmith present with his little collection of silver

utensils and ornaments of various kinds. Most of the metal he obtains

from miners of the same locality.

MARSTERS, PHYSIOGRAPHY OF THE PERUVIAN ANDES 257

Passing over the lowland of Cerro de Pasco, we come to the noted

deposit of copper-silver ore now in part the property of the Cerro de

Pasco Mining Company. The ore-body appears to be on or near the con-

tact of eruptives with a thick series of limestones. According to late

statistics, the amount of silver taken out of the surface portion of the

Cerro de Pasco deposit between the time of its discovery (1630) and the

end of the nineteenth century amounts to some 450,000,000 ounces.

If we should return to the coast from Cerro de Pasco via the Cerro de

Pasco Railway to Oroya, and thence via the Central Line to Lima and

Callao, we should pass through other mining localities such as Rio Blanca

and Morococha. Here again the respective ore-bodies are closely asso-

ciated with contact phenomena such as have already been described. In

the region of Matacana, a repetition of the same sort of occurrence may be

seen. In the region of Lima, there are copper-bearing ores associated

with eruptives in contact with limestones and shales. That is to say, the

occurrences of ores in each section can be correlated both geographically

and geologically.

Let us look for a moment at a south section across the Cordillera, say

from Mollendo to Puno and the east. At Mollendo, we find that outliers

of the Cordillera, or Cadena de la Costa, are composed in part of gneissic

and granitoid masses, which are probably intrusive in sandstones and

shales. Within the gneissic zones occur small copper deposits. While

an attempt has been made to develop some of these, they have never

reached the productive stage. Passing into the edge of the foothills in

the region of Carabaya, just below Arequipa, we again find a band of

erystallines bordered on each side by sandstones and shales. Prospect-

ing on a small scale has brought to light small bodies of copper-silver

ore, mostly located in the crystallines, while the sandstones and shales

are reported to carry coal beds. It is not until we reach Lagunillas that

we again come to a region containing ore deposits. Here are silver-

gold-copper ores in a belt extending from Santa Lucia to Marivillas.

This is to be correlated with the Caylloma silver occurrence already re-

ferred to. Here, as noted in other sections, we have eruptives associated

with limestones and shales. These are probably to be correlated with the

Morrococha Belt on the Lima section.

In the Titicaca basin to the south of Puno, ore deposits carrying silver

and lead have been prospected. I have not seen these, but I am informed

that the formations involved are the expected limestones associated with

eruptives. Geographically this should be correlated with the Cerro de

Pasco occurrence, as it lies in the Intercordilleran Belt and possesses

the same lithological relationships.

258 ANNALS NEW YORK ACADEMY OF SCIENCES

Within the Titicaca basin, we have an additional occurrence which so

far is not known to occur anywhere in the northwest extension of the

Intercordilleran Belt. In this basin, there is a considerable series of

sediments deposited upon lhmestones which are regarded as Cretaceous.

The former are probably Tertiary. They contain petroleum, but devel-

opment work has not been carried far enough to determine the areal extent

of the oil sands. The only work of any moment has been done by the

Titicaca Oil Company, backed by California people. Work has been

suspended for the present.

The gold-bearing veins of Poti as well as those of the Santo Domingo

region to the east from Tiripata are associated with the eruptives and

older sedimentaries of the East Cordillera. :

It is also known that the region of Huanaco, to the north of Cerro de

Pasco, is highly mineralized. From data at hand it would seem that the

geological associations are similar to those of Santo Domingo.

In the region of Huaraz and Recuay (valley of the Huaraz) there

exists another mineralized belt containing silver, copper and gold ores,

as well as lead ores. It is quite undeveloped. These are associated with

intrusives of the West Cordillera.

Iron, copper and silver ores also occur in the West Cordillera, to the

northeast of Piura. They are associated with dioritic intrusions and

bosses in the midst of a heavy series of shales and sandstones.

‘

RESUME

In the Andes of Peru, we can easily recognize a series of parallel and

well-defined physiographic provinces, which in the larger sense are defi-

nitely related to the geological development of the Andean Range as a

whole. .

Whatever has been the succession of physiographic changes in the

development of the Andes as a unit, there has evidently taken place, at

least on broad lines, a marked adjustment of drainage to structure, thus

affording a longitudinal and transverse arrangement, or pattern, such as

may be easily recognized in many other continental mountain systems.

A comparison of observed and recorded facts with reference to the

occurrence of ore-bodies in Peru proves that they are generally associated

with contact phenomena.

ns of the » Acalemy consist of two series, viz.

» mniform es a eee dollars oe volume. The articles

‘The ees of

aoe upon ee length and ie shies of illus-

ations, and may be learned upon application to the Librarian of the

c The author receives is ee as soon as his Daper has

tre a Fellows and ee a ‘The e

& Tue LIBRARIAN,

New York Academy of Sciences,

care of

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American Museum of Natural History, ;

New: York, Noe

NEW YORK

PUBLISHED BY THE ACADEMY A

to OctopER, O19:

Ae ae aitias 0

FES 6 101

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Naturat History, 1817-1876)

OFFicERs, 1912

President—EMERSON McMituin, 40 Wall Street

Vice-Presidents—J. EDMUND WoopMAN, FREDERIc A. Lucas

CHARLES LANE Poor, R. S. WoopwortH

Corresponding Secretary—Hunry EH. Crampton, American Museum

Recording Secretary—Epmunp Oris Hovey, American Museum

Treasurer—HENrRY L. DoHERTY, 60 Wall Street

Inbrarian—RaLPH W. Tower, American Museum

Editor—EDMUND Otis Hovey, American Museum

SECTION OF GHOLOGY AND MINERALOGY |

Chairman—J. E. WoopMaN, N. Y. University

Secretary—Cuag_Les P. Berkey Columbia University

SECTION OF BIOLOGY

Chairman—FREvDERIC A. Lucas, American Museum

Secretary—WiILL1AM K. GreGory, American Museum

SHCTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—Cuares LANE Poor, Columbia University

Secretary—F. M. Prprrsen, College of the City of New York

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. 8. WoopwortH, Columbia University

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The sessions of the Academy are held on Monday evenings at 8:15

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{ANNALS N. Y. AcAD, ScreNcES, Vol. XXII, pp 259-266. 15 October, 1912]

NOTES ON THE STRUCTURE AND GLACIATION OF

OVERLOOK MOUNTAIN

By Nein EK. Stevens?

(Read by title before the Academy 6 May, 1912)

CONTENTS

Page

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INTRODUCTION

Overlook Mountain is the southern terminus of the great central

mountain chain which forms the backbone of the whole Catskill system.

On the east, the mountain rises precipitously above the low ground of

the Hudson Valley, the land at the base of the steep incline on this side

being only about six hundred feet above sea level. This commanding

position gave Overlook for many years the reputation of being the high-

est peak in the Catskills, although it is actually more than a thousand

feet lower than Slide Mountain (Slide Mt. 4204 ft.; Overlook Mt. 3150

ft.). The position of Overlook in the Catskill system makes it of par-

ticular interest; and the present paper, though it by no means contains

a complete account of the geology of the mountain, is offered in the hope

that the notes contained therein may be of service to future investigators

and may, perhaps, stimulate an interest in the geology of this region.

The writer wishes to acknowledge his indebtedness to Professor H. H.

Gregory of Yale University for generous criticism and suggestions.

The summit of the mountain (Fig. 1) forms a triangle, from the

apexes of which project three main ridges with smaller ridges between.

The principal ridge stretches southwest for a distance of about four

miles and ends in a series of three lower peaks, separated from the main

peak by the Meads gap. The southern ridge is short and slopes rather

sharply to the level ground of the valley. The northeastern ridge, on

the other hand, is short and high, merging into the Plattekill Mountain

1 Introduced by James F. Kemp.

(259)

260 ANNALS NEW YORK ACADEMY OF SCIENCES

at an elevation of about 2500 feet. Aside from this narrow ridge,

Overlook is separated from the mountains to the north by the valley of

the Saw Kill. .

STRUCTURE

Unlike most mountains, the Catskills consist of a succession of piled

up, nearly horizontal strata, showing that they are really but the remains

of a great interior plateau now greatly eroded and dissected by water.?

VA aS IL Fa

w= —

SS = PGE

SE SFA

Fie. 1.—Overlook Mountain and adjacent territory. A portion of Kaaterskill Quad-

rangle, New York

U. 8S. G. 8. Topographic Sheet. Reduced. Scale: 1/83332

Except for a layer of hard conglomerate which caps many of the higher

peaks, the mountains consist almost entirely of alternate layers of soft

red shale and harder sandstone. The sandstone is sometimes thinly

laminated and frequently cross-bedded ; often however it shows a very even

2RALPH STOCKMAN TARR: The physical Geography of N. Y. State. New York, 1902,

p. 41.

STEVENS, GLACIATION OF OVERLOOK. MOUNTAIN 961

texture and is practically free from evi-

dence of fine stratification. This latter

variety of sandstone, generally designated

as gray or blue flagstone, is extensively

quarried for paving stone.

This alternation of soft, easily eroded,

shales with the more resistant sandstone

gives rise to the abrupt ledges, flat moun-

tain tops and “terraced” sides so character-

istic of the Catskills. Wherever a layer of

shale has been exposed, the surface has

been quickly eroded down to the next layer

of sandstone. Nowhere are these features

better shown than on Overlook Mountain.

Note in Fig. 1 the plateau-like tops of the

lower peaks and the terraces of the south-

ern ridges.

Fig. 2 shows a section through the two

lower peaks, between the Mead’s Gap and

the Bear Clove. This section extends from

where the strata first appear above the gla-

cial soil of the Woodstock valley, altitude

660 feet, to their summits, altitude 2100

feet. It is altogether probable that some

of the strata, represented here as of uni-

form thickness, are really somewhat lense-

shaped. ‘This however could not be deter-

mined owing to the prevalence of glacial

deposits. As will be seen from the figure,

the cap of these peaks consists of nearly

600 feet of sandstone. The upper portion

of this cap is hard and rather coarse, but

the stone becomes softer and more finely

lamimnated below. The lower layers of

sandstone are characterized by a _ cross-

bedded structure and contain numerous

streaks of red shale too thin to indicate in

the section. Besides the layers of sand-

stone and red shale there are, as indicated

by the figure, two layers of bluestone; the

upper outcrops at an altitude of about 1500

weet

er ey Cate

= O°

67 ONS . oO

. away te:

ON OOD Gyo ioya et Reh ls. 1°

° . es

Qe onOh eats

—————————————

460°

Shale = Bluestone WN

Covered] _|Sandstone|:2=~*.]

Fic. 2. —Diagrammatic section

through the two peaks west of

Meads Gap |

262 ANNALS NEW YORK ACADEMY OF SCIENCES

feet, the lower, which outcrops on the southern slope of the mountain,

is in this region covered with glacial drift. The strata as a whole dip

gently west-northwest.

GLACIATION

Two glacier streams have swept over this region: the Continental

Glacier, and later the more shallow Hudson Valley Glacier. The geo-

logic structure of the region makes it somewhat difficult to trace satis-

Fic. 3.—Map of Overlook region, showing direction of glacial strie (indicated by arrows)

Contour interval 200 feet

factorily the course of the glaciers; for the sandstones and soft shales of

which the mountain is composed have retained glacial markings only in

exceptional localities; while a considerable portion of the surface is, of

course, covered with glacial drift.

The map (Fig. 3) shows the direction of the striations left by the

Hudson Valley glacier in the Overlook region. It affords an interesting

3A, A. JULIEN: “The Excavation of the bed of the Kaaterskill.”” Trans. N. Y. Acad.

Sci., Vol. 1, p. 24-27. 1881.

STEVENS, GLACIATION OF OVERLOOK MOUNTAIN 263

example of the extent to which the direction of the ice near the surface

can be affected by the topography. A variation of more than ninety

degrees in the direction of the motion of the ice is indicated by the strize

on Overlook itself. Just northeast of Overlook, the course of the glacier

was nearly south (8. 20° W.). A portion of it, however, flowed over the

ridge between the Plattekill and Overlook Mountains and through what

is now the upper valley of the Saw Kill in an almost westerly direction,

moving nearly southward again through Meads Gap and at Shady. The

ice moved almost directly west up the Woodstock Valley, its course be-

coming gradually more southerly as it passed over the group of low hills,

known as the Ohio Mountain,* and the region farther south (S. 60° W.

near Glenford and 8. 40° W. at West Hurley).

MORAINES

While glacial action has probably not greatly altered the general out-

lines of the mountain, the valleys have been more or less filled with gla-

cial deposits. A moraine nearly a mile long fills much of what was once

the much deeper valley south of Meads Gap; while the upper valley of

the Saw Kill, between Overlook and the mountain ridge to the north, is:

filled to a considerable depth with morainic material. Both of these are,

in reality, parts of a large moraine which extends westward from Over-.

look.

As the moraines of the Catskills have been but little studied, a brief

account of the material composing them may be of interest. About:

eighty per cent of it is of local origin, consisting of the sandstone, shale

and conglomerate found throughout the Catskills. Of these materials,

conglomerate is the least common and forms less than ten per cent of the

whole. About one-half the local material consists of bowlders of various

sizes, with which pebbles and gravel are mixed with no sign of stratifica-

tion. From this it is apparent that water has played no part in the

deposition.

The foreign material consists largely of quartz and several kinds of

granite, with occasional pieces of water-worn, stratified rock and some

sandstone containing brachiopodous shells. Some shells picked up in

the bed of the Saw Kill, about two miles from its source, have been iden-

tified as Spirifer arrectus,® a species characteristic of the Oriskany sand-

4This mountain is called ‘“‘Tontshi Mt.” on the U. S. G. S. topographic sheet. It

seems, however, that this must be an error, as this elevation is locally known only as

“Ohio Mt.”: while the name “Tontshi” is applied to the much higher peak, left un-

named on the government map, just east of Ticetonik.

5The writer is indebted to Professor Charles Schuchert of Yale University for the

identification of this specimen.

264 ANNALS NEW YORK ACADEMY OF SCIENCES

stone, a thin layer of which outcrops in the Little Catskills near the

Hudson River due east from Overlook Mountain. It also appears in the

Helderbergs to the northeast.

One rather unusual variety of metamorphic rock, several pieces of

which were found, has been kindly identified by Professor J. F. Kemp

of Columbia University, who writes: “It is a type of rock fairly well

known in the Adirondacks. It has obviously been pretty well crushed

and granulated, but it is a member of the anorthosite series, which when

unchanged has large rectangular crystals of labradorite in a mass of

small crystals of augite. . . . The rock outcrops at the very head-

waters of the Schroon River in North Hudson township and also in the

Keene Valley. I think it probably occurs in many other places, where

it has not yet been specially observed. . . . In Bulletin No. 138 of

the New York State Museum, on page 43, under the name of “The New

Pond Locality,’ you will find a brief description of the rock.”

That both the specimens just referred to are characteristic of the Adi-

rondacks, together with the fact that all the metamorphic rocks found

are common in those mountains, indicates that most of the foreign ma-

terial in these moraines is of Adirondack origin.®

DRAINAGE

Except for a small portion of its eastern slope, Overlook is drained

entirely by a single stream, the Saw Kull, which forms a loop, some ten

miles or more in length, extending nearly around the mountain (Fig. 1).

From its source in Echo Lake at the base of the northeastern ridge, it

flows directly southwest through Shady, then east through the Wood-

stock Valley to a point directly south of its source. In the course of

this loop, the Saw Kill receives the smaller streams which flow from both

sides of the Overlook ridge. Owing to the small size of their watersheds,

many of these smaller streams are dry during a part of the year. The

rainfall is much greater in the spring than at any other season, and this

gives rise to floods which make the erosive power of these streams much

greater than it would otherwise be. The floods at this time are greatly

increased by the melting of the winter’s snow, and there is added the

erosive force of the ice as it breaks up.

6 These moraines yielded more foreign bowlders than did the ones on the north side

of the district, in the valley of Schoharie Creek, described by J. L. Rich in the Jouwr-

nal of Geology, vol. 14, p. 113, 1906, especially p. 120. Mr. Rich found but one bowlder

of gneiss. It may be that later glacial action, radiating from local centers, had con-

cealed earlier bowlders, brought in from the north. Mr. Rich’s paper has also some

general comments on the movement of the continental glacier, and, at the outset, upon

the present conditions of rainfall.

STEVENS, GLACIATION OF OVERLOOK MOUNTAIN 265

Although no data as to the actual stream-flow on Overlook are at hand,

a comparison of the flow of neighboring watersheds can not fail to be of

interest in this connection. Fig. 4 gives the comparative discharge of

the Schoharie, Esopus and Catskill Creeks for the different months

over a period of years.‘ It shows that more than one-third of the total

run-off of these streams occurs during two months, March and April. As

the curves represent an average of several years, they give but little idea

of the size of some of the floods. In the Schoharie, for instance, the

maximum daily discharge in November, 1907, was 13,100 cubic feet per

Cyonio =

SOPs: -———

SCHOHARIE =~

Wie. 4.—Average discharge of the Catskill, Esopus and Schoharie Creeks for each month

over a period of years

Expressed in second-feet per square mile run-off. Taken as follows: Catskill Creek,

South Cairo, N. Y., for period 1901 to 1905; Esopus Creek, Kingston, N. Y., for period

1901 to 1906; Schoharie Creek, Prattsville, N. Y., for period 1905 to 1908. Data com-

piled from Summary of the Climatological Data for the United States by sections. Sec-

tion 104 (Weather Bureau) ; Water Supply and Irrigation Papers Nos. 166 and 202;

and the Report of the State Hngineer and Surveyor, State of New York, 1907 and 1908.

second, while for the same month, in 1908, the maximum was onlv 268

cubic feet per second.

The valley of the Saw Kill is primarily of erosive origin. Like many

7 Records of rainfall, kept at Reservoir No. 1 of the Kingston City Water Works, in-

dicate that the rainfall near Overlook is much like that of the other watersheds men-

tioned.

266 ANNALS NEW YORK ACADEMY OF SCIENCES

other valleys in the Catskills,* however, it was partly filled with debris

by the continental glacier, so that now the Saw Kill, for the first five

miles of its course, has cut its way through a moraine of varying height,

70 ft. at Shady, and 40 ft. half a mile below its source. This fact is

clearly shown by the character of its bed, which is strewn with huge

bowlders of bluestone, sandstone and conglomerate, together with smaller

ones of granite and quartz, the harder ones still showing the marks of ~

glacial action.

Ecuo LAKE®

Echo Lake, the only considerable body of water near Overlook, is

clearly of glacial origin. It is a shallow pond, about three hundred

yards long by two hundred wide, and about eighteen feet deep in the

deepest part, situated in the angle formed by the Plattekill and Overlook

Mountains, just at the base of the high ridge connecting them. Across

this deep valley, the moraine forms a huge natural dam which holds

back the water of Echo Lake. The lake is thus bordered on three sides _

by high wooded ridges; while on the west, extending out over the mo-

raine, is a swamp larger than the lake itself.

As is to be expected from its situation, Echo Lake is apparently de-

creasing in size rather rapidly. The swamp on its westeru side is slowly

invading its waters. On this side, too, the lake is being narrowed by

the action of its outlet, the Saw Kill, in cutting back through the Glacial

drift which forms its bed. The pitch of the Saw Kill, which falls 1200

feet in the first four miles of its course, together with the floods men-

tioned above, makes this erosion relatively rapid.

In addition to this cutting away on its lower side, the lake is being

rapidly filled in from above. The silt and leaf mold washed from the

steep mountain ridges above the lake are deposited in the still water, and

the amount of this material is very considerable. On the north and east

this deposit forms a bed extending into the lake for more than two hun-

dred feet and reaching a depth of four or five feet. Here the deposit is,

to a considerable extent, protected from further action of the water by a

dense growth of the yellow pond lily, Nymphea advena, for which the

fine silt and leaf mold furnish a favorable substratum. The combined

effect of these agencies in reducing the size of the lake is so great as to

make it probable that, at no very distant date, Echo Lake will be oblit-

erated.

WasuHineron, D.C. —

8 Joun C. Smock: “On the Surface Limit or Thickness of the Continental Glacier in

New Jersey and Adjacent States.” Am. Jour. Sci., vol. 25, pp. 339-350. 1883.

® Also known as Sheu’s Lake.

ons and oe of researches, ae ‘with the Tec-

s and similar matter.

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agaeiiean Museum of Natural History,

New Yorks oN.

PEDAL LOCOMOTION :

“AND OF THE

OF THE LIMBS IN HOOFED ANIMALS |

inn1aM K. Gregory

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Natura History, 1817-1876)

OFFICERS, 1912

President—EmeErson McMiuuin, 40 Wall Street

Vice-Presidents—J. EDMUND WoopMAN, F'REDERIC A. Lucas

CHARLES LANE Poor, R. 8. WooDwoRTH

Corresponding Secretary—Henry HK. Crampton, American Museum

_ Recording Secretary—Epmunp Oris Hovey, American Museum

Treasurer—HeEnRY L. DoHzERTY, 60 Wall Street

Iibrarian—RauPu W. Towser, American Museum

Editor—EpmunpD Otis Hovny, American Museum

SHCTION OF GEOLOGY AND MINERALOGY

Chairman—J. E. WoopMAN, N. Y. University

Secretary—Cuartes P. Berkey Columbia University

SECTION OF BIOLOGY

Chairman—FREpDERIC A. Lucas, American Museum

Secretary—WiLuL1amM K. Grecory, American Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—Cuar_Es LANE Poor, Columbia University

Secretary—F. M. PEDERSEN, College of the City of New York

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. S. WoopwortH, Columbia University

Secretary—FREDERIC LiyMAN WELLS, Columbia University

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. Acad. ScrENCES, Vol. XXII, pp. 267-294, Pl. XXXIV.

18 October, 1912]

NOTES ON THE PRINCIPLES OF QUADRUPEDAL LOCOMO-

TION AND ON THE MECHANISM OF THE

LIMBS IN HOOFED ANIMALS

By WILLIAM K. GREGORY

CONTENTS

Page

NATTA CECE [N@ Inmet ameen ter Tepe Aca apes screen, cls meg eivariclrs lar WleGieca! ai bud) Auoie Water aly srisice Boag leks 268

PUMP SeEESALCeMa se COMMPOWMIG NEVES! tn cece cede ais celsleelc weiss veces cee 269

Hvolutionary stages of quadrupedal locomotion..................+.++---- 270

Factors of long distance travelling power in ungulates.................. 270

TENG NTO ORS Si Seen oe eee ee BS 6 ot Os RC RI eae 271

Adaptations for minimizing waste of energy..............+-+-.eee0: PAT

IEGMTRETITITT. & Soci etree ae RISO Bee cee ee ee ar a ee ea 271

Rropulsionyand= thexcenter Of Gravity. . ss. 0s5o-s - scr s+ eee ke oe eos 272

Sinuous movement of the body in running........................ 272

Spiral configuration of limb bones and articular surfaces.......... 273

AO Seamer Oil OMe eee rete ees aot A sere cates ayo icra Sieud Sie jes suswaieod! » aUolsterwheu a ate ae 273

Renaiihummrerionuota the Mim See = ice sides cad as cee ea aee wo oie alte sin Sere 273

SYOESGle coreicer. do G24 ca luteic sen calc ARERR SkG MCMC cl asset ok cr ares ike eo pee a 273

SGN Oe titel CL Caeewayey en eya actrees caer eoevede cue soe. ale fe oie ecu eb eis se rue gue e le Hee Siere 274.

eT SsalaO fannie raee oa remit an reetn Ala eos lie Cale ee ba ie cles ae 274

ANOGUE Ol -SUIBIGIO SSG Scie oo GcOte SRaraNe GOREN, ee een Rone a Pe OTE Pi ee 274.

Acceleration increment of stride due to ballistic power of limbs.... 274

HFECHeA TON Mit yan Clgn SUT G] Comers ieee evo sia cielerts elalei's eaeieis cece; ay aveney ) cneusevcis suse erates eye epaue 276.

Mechanical and physiological relations of “power” and “speed” in the loco-

SOTVOUEL WE, OVO ICTUIS 5 ceo ele ee Bec ye ous SG ser UE en OS aoa ea eT!

Conitrachle toreevor locomotive muscles: <2 5.46 fadeansc. ce os cess eee ces 276

Augmented by “hold and let go” arrangements................... DT

PAM OUT sy Olen OMG AGT O Mars tercnstar ys doll MAN Face SSE udtlagc, wage ravens Rake a eva ele PST

The “angle of insertion” and the principle of the parallelogram of forces. 277

Relations of speed of movement and angle of insertion............ 279

Variableness of the rotation component......................i... 219

“Moment of Resistance” and “Diminishing Load”..................... 280

Summary of mechanical and psychic factors in power and speed........ 287

Application of the foregoing principles to the study of the limbs of ungu-

VAS Seer teaerer ew cree epee Laie rata aye co au Ne iraafeurer aya been chase Chonan ay ene aaolo eon aconaeica gee oer 282

Functional significance of the angulation of the limbs................. 282

Mechanics of the foot in graviportal and cursorial types.>............. 283

MerAarSO-KEMVOLal Tm CAlOS ace oieie trie se erelc wtciete aia clejeed sie alae snes 284

Otper-adaptuve contrasts! im the feet. .3......5 625.052.2502. c i ee eee 287

Graviportal and cursorial types of tibia. ............... cee eee eee es 288

Cravaportalvand cursoniall types of femur. 2.25... seece. cee tee alee be 290

Graviportalandscursorial bypes Of Pelvis... 5)... . .ctslcs eee cee betes 292,

Graviportal and cursorial types of fore limb......................-.--- 293

Diagnostic vs. convergent evolution values of limb ratios.............. 294

Comparative: table of limb ratios. 22% 6... .6. 2b ote lee nee Facing page 294

(267)

268 ANNALS NEW YORK ACADEMY OF SOIENCES

INTRODUCTION

The movements and locomotive mechanisms of animals were the

subject of a classic work by Borelli in 1680, entitled “De Motu Ani-

malium.. .” The chief pioneers of modern research were the brothers

Weber (Eduard and Wilhelm), authors of “Die Mechanik der mensch-

lichen Gehwerkzeuge,” Gottingen, 1836. Marey, the author of the hand-

book on “Animal Mechanism”! (1874), invented elaborate apparatus for

analyzing and graphically recording the movements of the limbs of men

and animals, while more recently the mathematics of human locomotion

have been developed by O. Fischer and others. General reviews of animal

mechanics and of human locomotion are given by Marey,” Haycraft? and

Luciani. A very able analysis of the mechanism of locomotion in the

horse is given in “The Horse in Motion,” by J. D. Stillman,® while the

extensive series of photographs by Muybridge® record the actual positions

of the limbs and body assumed in motion by ungulates and other animals.

None of the above mentioned works considers the subject from the evo-

lutionary point of view. Ryder,” and especially Cope,® pointed out cer-

tain adaptations in the feet of ungulates, such as the reduction of the

digits in the “Diplarthra” and the so-called displacement of the metacar-

pals, and Cope used these observations in his argument for the hypothesis

of the transmission of acquired characters. He also made the following

very important observations :°

“In animals which leap, the distal segments of the limbs are elongated; in

those which do not leap, but which merely run or walk, it is the proximal seg-

ments of the limbs which are elongated.

“Animals which run by leaping are divided into those which run and leap

with all fours, as Diplarthra, and those which run and leap with the posterior

limbs only, as the jerboas and kangaroos. In both types, the distal segments

of the hind limbs are elongated, and in the Diplarthra, those of the fore limb

also. ale

“Animals which do not leap in progression (elephants, Quadrumana, bears)

are always plantigrade and have very short feet but elongate thighs and

mostly tibias.”

112mo. London, 1874.

2 Op. cit.

3H. A. SCHAFFER: Text book of Physiology. Edinburgh and London, pp. 228-273,

1900.

4Physiologie des Menschen . . . Ins Deutsche tibertragen and bearbeitet von Dr.

Silvestro Baglioni und Dr. Hans Winterstein . . . Siebente Lieferung. Jena, 1906.

5 Hxecuted and published under the auspices of Leland Stanford, 4to, Boston, 1882.

6 Animals in Motion . . . Third Impression 4to, London, 1907.

7Am. Nat., vol. 11, p. 607. 1877.

8“The Mechanical Causes of the Development of the Hard Parts of the Mammalia.”

Journ. of Morphology, vol. 3, pp. 137-290. 1889.

8 Tbid., p. 151.

GREGORY, QUADRUPEDAL LOCOMOTION 269

Cope neglected to follow up the important mechanical and adaptive

corollaries of these facts. He merely drew the very questionable infer-

ence that “those elements which receive the principal impact in progres-

sion are those which increase in length [in phylogeny ].”?°

In his paper on “The Angulation of the Limbs of Proboscidia, Dino-

cerata, and other Quadrupeds, in Adaptation to Weight,” Osborn? con-

cluded that

“The straightening of the limb [in the Dinocerata, Proboscidia, etc.] is an

adaptation designed to transmit the increasing weight through a vertical shaft.

Correlated with it are the shifting of the facets into the direct line of pressure

and the alteration of their planes from an oblique to a right or horizontal

angle with relation to the vertical shaft.”

Gaudry,’? in describing the limbs of extinct South American ungu-

lates, endeavored to show how the pose of these animals could be inferred

from a study of the facets——an idea which had been previously advanced

by Osborn.** Gaudry also designated as “rectigrade” the pose of ele-

phants and similar heavy forms which stand with straightened limbs and

toes, resting the weight chiefly on the pad.

The marked contrasts in the limbs and musculature between the slow-

moving heavy-bodied ungulates and the slender swift-footed or cursorial

types 1n various phyla, constitute a subject which will be discussed

in considerable detail in the monograph on the Titanotheres by Professor

Osborn. At his suggestion, the following notes, forming a part of the

present writer’s studies on this subject, are now published, together with

some of the drawings which have been made by Mr. Erwin 8. Christman,

under the direction of the writer, for the monograph above mentioned.

The writer is also indebted to Dr. W. D. Matthew for valuable criticisms

and suggestions.

LIMBS REGARDED AS CoMPOUND LEVERS

The simple principle that the limbs of quadrupeds are compound levers

and that the relative lengths of the upper, middle and lower segments ©

are adapted to specific loads, muscular powers and speeds, although well

understood by students of human and equine locomotion, has apparently

not hitherto been applied to elucidate the adaptive contrasts between

eursorial and “graviportal”’** ungulates.

10 Loc cit.

11 Am. Nat., vol. xxxiv, pp. 89-94. 1900.

12 ““Hossiles de Patagonie. Les Attitudes de quelques Animaux.’’ Ann. de Paléon-

tologie, tomei.

2 ‘‘Patriofelis and Oxyena restudied as Terrestrial Creodonts.” Bull. Amer. Mus. Nat.

Hist., Vol. 13, pp. 270, 271. 1900. :

144This word has been invented by Professor Osborn to describe the conditions in

heavy-bodied animals with long proximal and short distal limb segments.

Q"0 ANNALS NEW YORK ACADEMY OF SCIENCES

EVOLUTIONARY STAGES OF QUADRUPEDAL LOCOMOTION

In the Stegocephalian stage of quadrupedal locomotion, the short limbs

were held widely outward from the body, the humerus and femur were

very short and the feet were spreading and flat. Crawling was effected

in part by a sharp downward pull of a proximal segment (humerus or

femur), thus tilting the body upward on the same side and throwing the

weight on the opposite foot. The long axis of the body was meanwhile

thrown into alternate lateral curves, the advancing fore limb being on a

convexity, the advancing hind limb on a concavity.

In the late reptilian or early mammalian stage, the feet were brought

around partly under the body, the elbow and knee began to be drawn in,

the scapula was rotated backward as the coracoid lost its connection with

the sternum, and the body became well raised off the ground. According

to a hypothesis advanced elsewhere by the writer,*® this process was asso-

ciated with the acquisition of climbing, or semi-arboreal, habits, struc-

tural vestiges of which remain in the partly divergent first digit and many

other characters of early Eocene mammals."®

The Lower Eocene ancestors of the various orders of peedirtes had

probably all long since passed through these earlier stages of quadrupedal

terrestrial locomotion, and at that time many of them had perhaps become

more or less digitigrade. The primitive “Protungulates,” Meniscothervum,

Periptychus, Pantolambda, may give us some idea of what the several

ancestors of the subungulate series (Hyracoidea, Embrithopoda, Pro-.

boscidea, Amblypoda) may have been like. They also preserve apparent

traces of arboreal ancestry in the relatively short, spreading hands and

feet, long limb bones, the humerus with large entocondyle and entepicon-

dylar foramen, the undiminished power of pronation and supination of

the forearm and many other characters. The Basal Hocene Huproto-

gonia, the ancestor of the Condylarth Phenacodus, with slender subun-

guligrade feet, represents a more advanced stage of evolution, in the

direction of the Perissodactyls.

FACTORS OF LONG-DISTANCE TRAVELLING POWER IN UNGULATES

The primitive ungulates of the Lower Hocene were doubtless sur- .

rounded by environmental conditions which set the premium of survival

upon improvements in long-distance travelling power and in speed.

These improvements have been attained in various ways and in the most

15 ‘The Orders of Mammals,” Bull. Amer. Mus. Nat. Hist., Vol. 27, p. 226, 1910.

16 MATTHEW: Arboreal Ancestty of the Mammalia, Amer. Nat., Vol. 38, 1904, pp.

811-818.

GREGORY, QUADRUPEDAL LOCOMOTION Oral

diverse lines of evolution,—in the elephant no less than in the antelope.

The general factors of long-distance travelling power may be grouped

broadly under the headings: (A) Endurance and (B) Speed.

A. ENDURANCE

Endurance may be measured either by (1) the length of time an ungu-

late can keep in motion without rest or refreshment, or by (2) the.

amount of reserve strength left after a stated expenditure of energy, or

by (8) the relative quickness of recuperation. Endurance increases with

practice. As metabolism increases, the muscles, lungs, heart and other

organs of the thorax become stronger and larger.

More rapid metabolism requires more food and larger digestive ap-

paratus. Although the Lamarckian hypothesis in its crude form is very

probably untenable, it is a fact that herbivorous animals have longer and

heavier digestive tracts than carnivores. Moreover, as fast as the denti-

tion has become adapted for the harder, less nutritious kinds of food, the

digestive apparatus must have become more complex and wuch heavier.

While the enlarging thorax and abdomen have made availal.‘e a great

increase in energy, they have caused an even more rapid increase in total

weight.

With increasing total weight, the internal and external resistance to

be overcome in locomotion also rises, and hence the margin between the

total energy available and the energy required in progressing a given dis-

tance-is lessened. In other words, endurance, the first great factor of

long-distance travelling power, is directly proportional to the efficiency

of the adaptations for minimizing the waste of energy.

ADAPTATIONS FOR MINIMIZING WASTE OF ENERGY

Higher efficiency in locomotion has doubtless been attained, first, by

advantageous modifications of the organs of propulsion (such as are de-

scribed below), secondly, by improvements in the supporting frame-

work, thirdly, by improvements in the methods (a) of conserving the

inertia of forward motion, (0) of taking up shock, (¢) of preventing dis-

location and (d) of minimizing lost motion.

Momentum.—The shocks and strains to which the locomotive apparatus

is subject vary with the momentum of the body in motion. Hence as

momentum is the product of mass by velocity, the shocks and strains

experienced by heavy animals in rapid motion are very great, and devices

for lessening them become conspicuous.

Pee) ANNALS NEW YORK ACADEMY OF SCIENCES

Propulsion and the center of gravity—Perfect quadrupedal locomo-

tion,” says Stillman,*’ “requires uniform support to the center of gravity

(of the whole animal) and continuous propulsion by each extremity in

turn.” In walking and running, by the straightening or extension of the

limbs, the center of gravity is raised and thrown in advance of the centers

of support. The body thus falls forward, the center of gravity describing

a curve of greater or less convexity, the forward motion being accelerated

by the thrust of the propelling limb.

‘The first shock of the downward fall in the running horse is taken up

by the forwardly stretched and slightly bent hind limb (Fig. 1) placed

beneath or in advance of the center of gravity; the gradually stiffening

muscles of the thigh and back checking the downward momentum (Still-

man, p. 91). The rearing muscles thus come into play and serve to let

the fore part of the body down gently.

Dislocation of the fully extended forelimb in landing is prevented

partly by (a) the crutch-lke action of the limb itself (which is slung

from the converging fibers of the serratus magnus attached to the top of

the scapula) and by (0) the contraction of certain muscles of the shoul-

der, neck and back (Stillman, pp. 61, 62, e¢ seq.). ‘To these arrange-

ments and conditions, Stillman attributes the absence of the clavicles in

the horse.

The center of gravity in the smoothly trotting horse describes a rela-

tively flat trajectory, whereas in the “bounding” movement, or gallop, the

center of gravity ricochets and the trajectory consists of a series of

cycloids of marked convexity. This mode of locomotion, while very

rapid for short distances, is too wasteful for heavy-bodied animals, which

require a relatively flat trajectory and a maximum saving of inertia.

Sinuous movement of the body in running.—By the bending backward

of the pelvis, first on one side and then on the other, the thrusts of the

femora are brought more nearly into line with the anteroposterior axis,

while wrenching of the pelvis is prevented by the contraction of the

longissimus dorsi of the opposite side (Stillman, p. 36). By this means

also, the length of the stride is directly increased. The same sinuous

motion of the body is associated with the “figure-of-8” movement of the

limbs noticed by Pettigrew.*® .

In this connection may be noted also the devices for avoiding “interfer-

ence” of the limbs (e. g., “stifle action” of the iliacus, preventing the knee

from striking the abdomen; oblique trochlea of the astragalus carrying

the advancing foot around its fellow of the opposite side). Dislocation

17 Op. cit., p. 87.

18 Animal Locomotion, p. 39. 12mo. New York, 1874.

GREGORY, QUADRUPEDAL LOCOMOTION 273

is provided against not only by the ligaments, but also by the metapodial

keels, by the grooved trochlea of the astragalus, by the cnemial crest of

the tibia, ete.

Spiral configuration of limb bones and of articular surfaces—Good-

sir, Pettigrew*® and others have shown that the articular surfaces of the

elbow, ankle and calcaneo-astragalar joints are spirally warped surfaces

which act after the manner of screws. The limbs as a whole also are

twisted levers with the ridges and muscles arranged spirally. “This

arrangement,” says Pettigrew, “enables the higher animals to apply their

traveling surfaces to the media on which they are destined to operate at

any degree of obliquity so as to obtain a maximum of support or propul-

sion with a minimum of slip. If the traveling surfaces of animals did

not form screws structurally and functionally, they could neither seize

nor let go the fulera on which they act with the requisite rapidity to

secure speed, particularly in water and air.”

Lost motion—Lost motion through backward slipping of the foot

upon the ground is provided against in the horse by the form and details

of the hoof, and in the elephant by the plantar pads.

Pendulum action of the limbs.—The brothers Weber held that in rapid

locomotion the limbs swing freely as pendula, but Marey and later in-

vestigators, according to Luciani,” hold that the natural swing of the leg

is very largely damped and controlled by the flexor muscles. In favor of

the view that there is some measure of analogy to the pendulum, we ob--

serve that in the horse, the center of gravity of the limb, corresponding:

to the “bob” of a pendulum, is relatively proximal in position, and this:

is associated with rapid oscillation of the limb, whereas in the elephant,

the center of gravity of the limb is farther down the shaft, and here we

have a slower oscillation of the limb. It will be observed that while the

body is moving forward, the propelling limb is moving backward, and its

own backward momentum, due to weight alone and to the pull of the ex-

tensors, must be overcome by the forward pull of the flexor muscles and

by the forward pull of the body as a whole. Hence the heavier the limb,

the greater the force expended in overcoming and reversing the mo-

mentum of each limb at the end of each stride.

SPEED

The speed of a quadruped or biped in motion is measured by the

product of the length of the stride into the rapidity of the stride.

12 Animal Locomotion, pp. 23-24, 28, 29. 1874.

20 Op. cit., p. 126.

274 ANNALS NEW YORK ACADEMY OF SCIENCES

The diverse adaptations in the limbs, considered as compound levers,

are related to either or both of the factors, “length of stride” and “rapid-

ity of stride.”

LENGTH OF STRIDE

Length of limb.—Length of limb is the first factor of length of stride.

Tt is generally proportional to height at the shoulders and hips. Length

of limb has been attained in cursorial animals by lengthening the lower

and middle segments of the limb, in graviportal animals by lengthening

especially the proximal segments of the limb.

Angle of stride—Angle of stride is the second factor. It is measured

by the are described by the lower end of the femur or humerus in swing-

ing from the position of extreme extension to that of extreme flexion. A

wide angle of stride not only lengthens the stride, but also enables each

limb, first, to be placed in turn below or beyond the center of gravity in

order to secure more continuous support for the center of gravity. and,

secondly, it enables the propelling limb to exert its propulsive effort for

a relatively long period. ;

Acceleration increment of stride due to ballistic power of limbs.—Those

portions of the stride that are due simply to the length of the limb and to

the angle of the stride might, if determined, be illustrated by moving the

inert limbs of a dead animal suspended in the air. In the slow walk of

a biped, the successive positions of the legs might for our purposes be rep-

resented by a series of inverted V’s (/\/\/\/\/\/\ ) with the lower ends

touching. Each /\ represents a single step and two successive /\’s repre-

sent a stride. In the rapidly moving animal, however, the stride receives

a very considerable increment, due to the impetus imparted by the pro-

pelling limb and to the forward motion of the body as a whole, which

carries the forwardly moving foot to a position far in advance of its own

unaided reach. This “acceleration increment,” as it may be called, in-

creases with the velocity of the movement and is proportional to what

we may designate the ballistic power of the limb. This ballistic power

may be defined as excess propulsive power over and above that which

is necessary to move the limbs as stilts and to support the weight of the

body; it is expended in lengthening the stride. Ballistic power and

the acceleration increment of the stride are measured by the length of

time at least three of the feet are off the ground together during a single

stride of a quadruped running at full speed. In Fig. 1, representing an

elephant in rapid motion (ambling), it will be observed that three of the

feet are never off the ground at the same instant; whereas Stillman’s

Figs. 2-10 show a galloping horse in which at least three of the feet are

GREGORY, QUADRUPEDAL LOCOMOTION 275

off the ground at once in seven out of nine phases of one stride. The

same figures show that the elephant has three feet on the ground together

during five-sixths of one stride, and during the remaining sixth the body

is supported by one fore foot and the opposite hind foot, whereas the

horse in question never has three feet on the ground together during the

stride there pictured. Thus it will be seen that the “acceleration incre-

Fic. 1.—Graviportal and cursorial modes of locomotion contrasted in the amble of the

elephant (I) and gallop of the horse (II)

I after Muybridge; II after Stillman

ment” of the stride and the ballistic power of the limbs are at a maximum

in cursorial animals and at a minimum in graviportal animals. The

“acceleration increment” will no doubt increase also with the potency of

the psychic motive and of the neural stimulus (cf., p. 282).

In brief, the factors of length of stride are (1) length of limb, (2)

angle of stride, (3) acceleration increment.

2"6 ANNALS NEW YORK ACADEMY OF SCIENCES

RAPIDITY OF STRIDE

Rapidity of stride, the second major factor of speed, is determined by

the rate of oscillation of the limbs, especially of the proximal segments.

The conditions determining rapidity of stride are discussed below (p.

281).

MECHANICAL AND PHYSIOLOGICAL RELATIONS OF POWER AND SPEED IN

THE LOCOMOTIVE APPARATUS

CONTRACTILE FORCE OF LOCOMOTIVE MUSCLES

The contractile force of a muscle (1. ¢., its ability to overcome inertia

at a given instant) is proportional to the number of its contractile fibers,

when these fibers are parallel to the direction of contraction. 'The force

of such a muscle is therefore proportional to the sectional area of the

muscle.*t In the case of weights lifted vertically by isolated muscles,

the work (W) performed is measured by the product of the muscular

force (F), multiplied by the distance (D) through which the load is

hifted?? (W=F*X D). This distance is proportional to the length of

the muscle,* for the “shortening” of a muscle is proportional to its

length. Hence the total work performed will be proportional both to the

length of the muscle and to its sectional area, and hence to the mass or

the weight of the muscle.?*

The work performed by a long muscle is greater than that of a shorter

one of the same sectional area.2> Long and slender muscles such as the

sterno-mastoid and the sartorius of man exert a small power over a long

range; short and thick muscles such as the pectoralis major, the gluteeus

maximus or the temporalis develop a relatively great power multiplied by

a short range.2® The contractile force of muscle per unit of sectional

area is much less in cold-blooded than in warm-blooded animals. It is

lessened by disuse and extreme fatigue and is increased by exercise, and

hence is dependent upon the nervous system and general systemic condi-

tions.

The contractile force is inversely proportional to the number of con-

nective tissue fibers mingled with the striped muscle fibers. Hence

muscles grade into tendons and ligaments. When a muscle is stretched,

21 HAYCRAFT in Schifer’s Text hook of Physiology, p. 242.

22 Toid., p. 245.

23 Toid., p. 244.

24 Tbid., p. 246; also Marey, p. 62. 1874.

2 FAAYCRAFT, p. 246.

26 MARRY, p. 62.

27 HLAYCRAFT, p. 243.

GREGORY, QUADRUPEDAL LOCOMOTION any

it serves partly as a ligament. All muscles in situ are stretched to a

certain degree, and thus act as ligaments.** “Over extension” of the

muscle is prevented by the inextensible connective tissue fibers.*° Ac-

cording to Stillman,*° the length of the muscles cannot be increased by

exercise, otherwise the tension necessary to prompt action would be lost.

The contractile force is highest when a muscle is stretched to its full

“physiological length” (that is the greatest length it ever assumes during

life). As shortening takes place, the contractile force becomes less and

less (Haycraft) .**

Contractile force and speed of movement augmented by “hold and

let go” arrangements.—Fick and Helmholtz showed** that the greatest

force and velocity of contraction are developed when the movement of the

muscle is checked during the initial stages and when the resistance is

suddenly diminished.

Amount of contraction.—The shortening of individual muscles is in

general proportional to their length when in repose, but different in-

vestigators give somewhat different estimates. “While Weber described

a muscle as shortening 70 per cent. of its length, when unweighted, more

recent observers incline to put the shortening at 20 to 30 per cent. of its

length” (Haycraft).** Marey** estimates “the mean shortening of a

muscle while contracting, when it is not detached from the animal,” as

“about a third of its length when in repose.” Bishop’s estimate is one-

fourth (Stillman, p. 31). “When the fibers are not parallel, but

obliquely set, as in the gastrocnemius, we have a greatly extended trans-

verse area of muscular fibers, which act therefore very powerfully,

though, on account of their short length, they can exercise their pull but

a comparatively short distance” (Haycraft) .*°

THE “ANGLE OF INSERTION” AND THE PRINCIPLE OF THE PARALLELOGRAM

OF FORCES

In Fig. 2 (1), let AC represent a rod free to rotate around the point

A in the direction CC’; let BD represent a contractile spring fastened

at D, inserted on AC at B and forming the angle ABD (a). Assume

that the length of BD is proportional to its contractile force; then from

28 Tbid., p. 245.

23 Toid., p. 242.

30 Op. cit., p. 32.

31 Op. cit., p. 242.

34 HAYCRAFT, op. cit., p. 248.

33 Op. cit., p. 244.

34 Op. cit., p. 62.

33 Op. cit., p. 242.

278 ANNALS NEW YORK ACADEMY OF SCIENCES

the principle of the parallelogram of forces, we may resolve BD into two

forces, the first AB acting in the direction of the rod AC and tending

to press AC against its fulerum A, the second component BR acting at

right angles to the first and tangent to the are of rotation BB’. The

first component AB may be called the “centripetal component,” the sec-

ond BR may be called the “rotation component.” In Fig. 2 (IL), the

contractile spring bd is of the same length as before, but the angle of

“i

Fic. 2.—Diagram illustrating the direct relation of the angle of insertion (a,a’) to the

“rotation component” (BR, br) and the inverse relation of the angle of inser-

tion (a, a’) to the “centripetal component’ (AB, ab) and to the speed of the

insertion point (proportional to BB’, bb’).

insertion abd (a’) is increased; then the centripetal component ab will

be less than AB, but the “rotation component” br will be greater than

BR. Accordingly as the angle of insertion increases, the pull across the

shaft becomes more direct, while the pull along the shaft decreases; in

other words, the rotation component varies directly, the centripetal com-

ponent inversely, with the angle of insertion.

GREGORY, QUADRUPEDAL LOCOMOTION 349

Applying this to Fig. 6, Il, we see that in the horse, the muscles fig-

ured are inserted at more open angles of insertion (a, B, y) than in the

mastodon or elephant and that their rotation components are therefore

relatively greater, the centripetal components relatively less.

The foregoing principles were worked out independently by the writer,

but Luciani*® gives similar principles for human locomotion. He states

that not all of the muscular force is available for the movement of the

skeleton, that this is only the case when the insertion of the muscle on

the bone reaches almost a right angle, as in the case of the masseters,

which can exert their whole strength in pressing the lower jaw against

the upper jaw. He says that generally, owing to the conditioning form

of the skeleton, the muscle is attached more or less obliquely, so that the

direction of its fibers makes a more or less acute angle with the long axis

of the bone. In all these cases, a great part of the total force of the

muscle is lost as regards movement. In every case, however, whatever

the form of the muscle or the size of the angle of insertion may be, by

resolving the total pull into its components, in accordance with the law

of the parallelogram of forces, one can determine how much of the total

pull is expended in the movement of the bone, assuming the other bones

to be stationary. Liuciani*’ also shows that the more acute the angle of

insertion is, the smaller will be the component of rotation, and the nearer

the angle of insertion approaches a aga angle, the greater will be the

ee of rotation. :

Fig. 2, we see that if DB contracts to DB’, the point of insertion will

move from B to B’. If now the angle of insertion be increased to a’ and

db (equal to DB) contracts to db’ (equal to DB’), then the point of in-

sertion moves only through bb’, which is less than BB’. If the contrac-

tion time as well as the distance be constant, then B will move faster

than b; that is, when the rate of contraction and length of muscle are

constant the speed of the insertion point varies inversely with the angle

of insertion.

It is also evident that if the angle of insertion and other factors re-

main constant the speed of the distal end of a long bone will increase as

the point of insertion is moved toward the head of the bone. (Because

BC will be larger.)

Variableness of the rotation component—From Fig. 2, it will be seen

that the angle ab’d is somewhat greater than abd, that is, both the angle

of insertion and the rotation component increase as the muscle contracts.

36 Physiologie des Menschen, Siebente Lieferung, p. 115.

31 Tbid., p 116.

280 ANNALS NEW YORK ACADEMY OF SCIENCES

“MOMENT OF RESISTANCE” AND “DIMINISHING LOAD”

In every lever, whether of the first, second or third order, the “power”

and the “resistance,” acting along parallel lines, but in opposite direc-

tions, are in equilibrium when the power multiplied by its effective dis-

tance from the fulcrum is equal to the resistance multiplied by its effect-

ive distance from the fulcrum. The “effective distance” is measured by

a line passing through the fulerum and perpendicular to the line of direc-

tion of the force. The product of a force multiplied by its effective

distance from the fulcrum is called ds “moment.”

M W

Wie. 3.—l. Hindfoot of an extremely cursorial type (Neohipparion) showing at the in-

stant of greatest extension of the foot a low moment of power of the calf

muscles (M X BB’) and a very high moment of resistance (W X B’ A’ v

of the pressure of the tibia upon the ankle.

II. Hindfoot of an extremely graviportal type (Mastodon) showing at the in-

stant of greatest extension of the foot a much higher moment of power of

the calf muscles (m X bb’) and a relatively lower moment of resistance

(w xX b’ a’) of the pressure of the tibia upon the ankle.

In Fig. 3, I, it would appear natural to assume that the point A, on

the ground, is the fulerum, and that the “resistance” is the pressure of

the tibia upon the ankle joint at B’, while the “power” is the contractile

force of the muscles of the calf, applied at B. Similarly, Eduard

Weber®® described the human foot as a lever of the second order and gave

for the relations of the forces and movements of the foot in raising the

33 Cf. HAYCRAFT, op. cit., p. 251.

GREGORY, QUADRUPEDAL LOCOMOTION 281

weight of the body a formula which translated into the terms of our

Fig. 3 would be as follows:

M X BA’=W X B’A’

But Knorz, Henke, Ewald and others, as quoted by Haycraft (Joc. cit.),

showed that the effective distance of the muscular force M is not BA’,

but BB’, and that we should rather conceive the foot as a lever of the

first order with the pivot at B’, the “power” at B and the “resistance”

(offered by the reaction of the ground upon the foot) at A. In that

ease, the moments around B’ are ag ollows :

M X BB’=W X B’A’

Hence, other things being equal, the longer the foot, the greater will

be the moment of resistance to be overcome by the muscles of the calf.

If the angle B’AD’ be increased, as when the foot assumes a more

vertical position, the effective distance B’A’ decreases; that is, the mo-

ment of resistance decreases as the foot becomes more vertical. In other

words, the “load” diminishes as the calf muscles contract. It has been

shown by Fick and others (quoted by Haycraft, loc. cit., p. 246) that

when the force of- muscular contraction is opposed to a diminishing mo-

ment of resistance, the muscle is capable of performing more total work

(force X distance) than when the resistance is constant. Consequently,

the diminishing resistance, conditioned by the raising of the foot, ena-

bles the calf muscles to perform their work under the most favorable

conditions.

SUMMARY OF MECHANICAL AND PSYCHIC FACTORS IN POWER AND SPEED

The speed of the distal end of a “long bone” of the limbs will depend

upon (1) the nearness of the point of insertion of the principal muscles

to the joint or axle, (2) the smallness of the angle of insertion of the

muscles, (3) the position of the muscle fibers with reference to the long

axis of the muscle, and (4) the speed of contraction of the muscle itself.

If a long muscle and a short muscle were isolated for experiment, it

might prove that the short muscle would contract faster than the long

one, but, in nature, a single movement of a long bone is produced by the

simultaneous and coordinated action of muscles of varying length. Thus,

in the act of extending the whole arm from the fully flexed position,

the relatively short, broad thoraco-scapular and scapulo-humeral muscles

contract in the same time as do the relatively long extensors of the fore-

arm, irrespective of their lengths. The speed and force of contraction

naturally depend partly upon the strength of the stimulus and partly on

989 ANNALS NEW YORK ACADEMY OF SCIENCES

the resistance to be overcome.*® The end result, as it were, determines

the rate of contraction of the codrdinated muscles. The effective regu-

lation and correlation of muscular action is obviously an extremely com-

plex function of the peripheral and central nervous systems. The force

and speed of contraction of a given set of muscles in a living animal at

a given moment are determined not only by many mechanical factors, of

which a few have been mentioned above, but also by the whole psychic

constitution of the animal and by the psychic effectiveness of the ex-

citing “motive.” .

Fic. 4.—‘Graviportal”’ adaptations for the walk and amble in the Mastodon

APPLICATION OF THE FOREGOING PRINCIPLES TO THE STUDY OF THE

Limps oF UNGULATES

~FUNCTIONAL SIGNIFICANCE OF THE ANGULATION OF THE LIMBS

As noted above (p. 269), the straightness of the limbs in the Probos-

cidea and similar heavy-bodied animals was interpreted by Osborn in

1900 as “an adaptation designed to transmit the increasing weight

through a vertical shaft.” While this is no doubt an incidental advan-

tage of the straightness of the limbs, it is probably not the chief teleo-

logical “object.” From a consideration of the mechanical principles

governing the use of the limbs as compound levers (see pp. 278-281) and

39 J. BuRDON SANDERSON, in Schiifer’s Text Book of Physiology, Vol. 2, p. 363, 1900.

GREGORY, QUADRUPEDAL LOCOMOTION 983

from a comparison of the photographs (Fig. 1) of an ambling ele-

phant and of a galloping horse, it seems probable that the straightness

of the limbs in graviportal animals has been evolved pari passu with the

short rectigrade feet and with an ambling even gait, in combination with

a long stride of minimal acceleration increment (p. 274). Conversely,

the bent or angulate character of the limbs in the horse and other cur-

sorial animals is correlated in part with the very long, slender unguli-

grade feet and with a bounding galloping or trotting gait, in combination

with a long, very rapid stride of maximal acceleration increment.

Fic. 5.—“Cursorial” adaptations for the run, gallop, etc., in the Neohipparion

In other words, the use and structure of the feet have been the teleo-

logical dominants which have determined the diverse modifications in

the musculature, proportions and angulation of the proximal segments

of the lhmbs, just as in early stages of aquatic adaptation in reptiles

(e. g., Thalattosuchia, Nothosauria, etc.) the aquatic habits are reflected

more clearly in the feet or distal segments rather than in the proximal

limb segments.

MECHANICS OF THE FOOT IN GRAVIPORTAL AND CURSORIAL FORMS

Comparing the structure and function of the graviportal and cursorial

types of feet, we see (Figs. 3 and 4) that, in the elephant, the very mas-

sive gastrocnemius and soleus muscles are attached at a wide angle to the

massive calcaneum, while the foot itself is very short. In the position

shown in Fig. 3 (11) and as compared with conditions in the horse, this

984. ANNALS NEW YORK ACADEMY OF SCIENCES

gives a relatively high moment of power (proportional to 6b’) to the

calf muscles and a relatively low “moment of resistance” (proportional

to b’a’) to the great pressure of the shaft of the limb upon the astrag-

alus. A considerable part of this “moment of resistance” is also sub-

tracted by the supporting effect of the great pad of elastic tissue under-

neath the foot. No doubt the specimen from which Fig. 3, II, was

drawn should have been mounted with the feet more nearly vertical ;

this would greatly shorten 6’a’ and further increase the advantage of

m X bb’. As the plantar pad is raised from the ground, more weight is

thrown on the toes, but, at the same time, they are brought further back

almost beneath the astragalus, thus reducing 6’a’ to a minimum, so that

the load decreases as the muscles contract. This arrangement not only

compensates for the fact that the greatest absolute force of a muscle is

developed when it is stretched to its full psysiological length, but it also

permits the muscle to perform a greater total quantity of work than

would be the case if the load were increasing instead of diminishing

(p. 280).

Similarly, in the horse, the greatest “moment of resistance” is when

the foot is fully flexed forward (which is at the instant the foot touches

the ground) ; the action of the extensor muscles is thus suddenly checked ;

this conditions physiologically a corresponding and sudden increase in

the available energy (p. 277). By the raising of the heel the moment

of resistance (W X BA’), as in the case of the elephant, also decreases,

a. @., the load diminishes; but in the elephant, the motion of the foot is

relatively slow and the acceleration increment of the stride (p. 275) is

therefore slight, whereas in the horse the motion of the foot is very

rapid, and the acceleration increment (through the high velocity im-

parted to the relatively light body by the spring-like extension or open-

ing of the angles at the stifle, hock, fetlock and pastern) is very great,

so much so that at least three of the feet are off the ground during a

great portion of the time. In brief, short feet (as in the Proboscidea)

slightly bending at the ankle, raise a heavy load through a short distance

with a minimal acceleration increment of the stride; long feet (as in

the horse), sharply bending at the ankle, throw a smaller load a long

distance, with a maximal acceleration increment.

Metatarso-femoral ratios.*°—That in cursorial animals the hind foot

is long as compared with the femur, while in graviportal animals the

40 The investigation of limb- and arch-form and proportions, and especially the estab-

lishment and significance of definite ratios between the limb segments, were suggested

by Professor Osborn and taken up conjointly by him and the writer in the Hocene sec-

tion of the Titanothere Monograph; the following observations on limb ratios are in

part a preliminary publication of the joint results attained.

GREGORY, QUADRUPEDAL LOCOMOTION 285

reverse is the case, is shown by the table of ratios, Plate XXXIV, and

especially in the following examples:

Length of metatarsal III

Pane: Length of femur

Graviportal Mediportal Subcursorial Cursorial

Coryphodon... .14 Rhinoceros .37 Eohippus... .50 Higquusisesaes 18

Uintatherium. .10 Paleeosyops .21 Tragulus.... .56 Antilope. ... 1.00

Mastodon..... met Mesohippus. .57 Odocoileus .. 1.00

Wlephas.... ....+. 13

Brontotherium .20

oxod om =). ....': Sly

These ratios definitely prove the connection between the mode of loco-

motion and the length of the middle metatarsal as compared with the

femur. The wide differences in the metatarso-femoral ratio, ranging

from .10 in extreme graviportal forms to 1.00 and upward in cursorial

forms, are partly bridged over in the mediportal and subcursorial types,

and even more completely in a fifth group including certain primitive

fee Mitse el:

ungulates, the Condylarths, in OS ae ranges from .43 to .31.

Perhaps the most important facts to keep in mind in comparing these

and similar ratios (below) are that the ancestral Placentals probably

had relatively short hands and feet and long limb bones (p. 270), but

that there was doubtless a considerable range of variation in this respect

even as far back as the Upper Cretaceous epoch. We are unfortunately

unable to follow the ratios through approximate phyletic series except

in a few cases (especially Titanotheres, Equid, Rhinocerotide), but, in

every case, we can feel sure that the precise ratios attained in the end-

forms are conditioned largely by the nature of the ratios in the stem-

forms of each family.

Thus, the exceptional shortness of the feet in the Amblypoda is con-

ditioned by the fact that this group, as represented by Pantolambda, had

comparatively short feet before gigantism was developed. Hyrax is

another example of a small form with very short feet, and from some

such forms the Proboscidea probably arose.

Besides those phyla which had short feet in the ancestral forms and

which merely emphasized this feature, there are many phyla which

started from animals with feet of moderate length and later shortened

up the feet to a considerable extent. Thus in the Titanotheres, the oldest

and most primitive form (Hotitanops) has a metatarso-femoral ratio of

about .34, which is not far from that of other early Perissodactyls, but

236 ANNALS NEW YORK ACADEMY OF SCIENCES

in the collateral descendants of Hotitanops, we observe a relative short-

ening of the digits, correlated with increasing body size and straighten-

: Mts. III.

ing of the knees, so that ce from .34 through .31 and .30

in the Middle Eocene genera to .28, .26 and even .20 in the gigantic

' Brontotherium. Again, in the Rhinoceroses, the oldest, smallest and most

ay Mts. IIT.

primitive forms have a foot of moderate length aod = 43-42),

but by progressive relative shortening and broadening, the ratio drops to

.24 in Metamynodon. In the Hippopotami, which are probably descended

from animals proportioned about as in Oreodon (with an index of .38).

gigantism and aquatic habits have brought about a reduction of the index

to .26. Similarly in the Toxodonts, the smaller and more primitive forms

had relative long, slender feet, while in the gigantic Toxodon

fallSto cai:

In those groups in which the most primitive known members had

already attained a slender foot, with reduced side toes, gigantism is

unable to effect a complete approximation to the graviportal type. Thus,

in the bisons, which are undoubtedly descended from slender-footed forms

having a metatarso-femoral ratio not less perhaps than .75, the sudden

increase in size causes the ratio to fall but slightly (to .65). In the.

gigantic Irish elk, whose ancestors probably had a very high metatarso-

femoral ratio (perhaps 1.00 or more), this ratio falls to .71.

The evolution of cursorial forms is also indicated by marked changes

in the ratio under consideration. Thus, in the Equide, it rises from

.53 in Hohippus, through .68 in Mesohippus to .99 in Hypohippus, reach-

ing the extreme of 1.16 in the slender-limbed Upper Miocene horse,

Neohipparion. In the relatively small and slender kiang, the ratio is

still 1.00, but in the relatively heavy-bodied Equus scotti of the Pleisto-

cene, the ratio is only .84, while in modern horses, we observe even in

race horses a falling off of the index to .78, and in the stocky-limbed

Hippidion, it drops to .72. Correlated with this fall in the length of

the metatarsal III, we observe a straightening of the knee. These fig-

ures possibly may mean that the modern Hquus caballus, on account of

its great size, 1s somewhat less adapted to extreme cursorial locomotion

than was the slender Neohipparion which closely paralleled the deer

Odocoileus.** On the other hand, race horses seem to have compara-

tively long femora (cf. Stillman, p. 80) and cheetahs, hounds and other

“J. W. GipLrey : Bull. Amer. Mus. Nat. Hist., Vol. 19, pp. 474-476. 1903.

Mts. ITI. -

GREGORY, QUADRUPEDAL LOCOMOTION 987

forms that progress by bounding have quite long femora. In this con-

nection, it must be remembered (p. 278) that a long femur, implying

small angles of insertion of the principal extensors, gives relatively high

speed of rotation of the insertion points, but low power for the rotation

components. ‘The long femur of graviportal forms has a wholly different

meaning (p. 289).

Progressive reduction of the side toes in the Artiodactyla as in the

Perissodactyla is accompanied by the elongation of the cannon bone

(here represented by two coalesced digits, metatarsals III. and IV.) and

by other cursorial adaptations. The primitive Artiodactyl foot with

four complete side toes is represented in Oreodon, which has a ratio of

.38. In Sus the ratio is .84, in the primitive four-toed camel Hotylopus

reedi it rises to .52, in Tragulus to .66, whence it rises rapidly to 1.00

and more in the deer and antelopes, culminating in 1.35, an extreme

figure, in the giraffe. In the giraffe, the development of great body

size has not brought about any reduction in the length of the metacar-

pals, but, in correlation with the long neck, has even lengthened them.

Other adaptive contrasts in the feet.—There are other adaptive con-

trasts in the feet of graviportal and cursorial animals, as follows:

In graviportal forms, the astragalus is flattened down, for, when the

weight of the body is raised, it is easier to force the tibia up a gentle

slope than a steep one. No lateral keels are needed on the trochlear

surface to prevent dislocation because of the breadth and spreading

character of the tarsus and of the large size of the fibular malleolus. In

eursorial animals, on the contrary, the curvature of the astragalar troch-

lear is steep and the range of movement wide; the trochlea keels help to

keep the narrow tarsus in place.

With regard to the phalanges, in cursorial animals, the ae bending

at the fetlock and pasterns and the sudden straightening out of these

joints under the pull of the powerful flexors of the foot greatly assists in

projecting the body into the air (Stillman, loc. cit., p. 89).

In graviportal animals, on the other hand, the terminal phalanges are

reduced ; the massive flexors of the foot raise the weight slowly and assist

the animal in rolling from one foot to the other.

288 ANNALS NEW YORK ACADEMY OF SCIENCES

GRAVIPORTAL AND CURSORIAL TYPES OF TIBIA

Some of the tibio-femoral ratios (=) given on Plate XXXIV may here

be grouped as follows: F

Graviportal Mediportal

intathertunie see eee 53 Palzeosyopshia yen... oe eee Teh

Cory phodom. cert sen eee cee 61 Rhinoceros indicus............. 79

LEV PONE MUI. Sy peoaueone dear sac 56 Tapirus.c) cs.6 22s. 2 eee .80

Mastodon rey an eee eee ner .69 Pantolambdas:.... 3,-2.5 .76

Hiephashndicusiy.eee eee .60

Brontotherium............. ... 54

Metamynodon.................. 08 Cir gorten

Peleoceras. eee ae cae .O7

: Eohippus \22.25 oes8 ao eee -. 1.00

Subcursorial or Primitive Mesohippuss).). 225m s4. 4-eeeeee 1.08

Phenacodus primeevus.......... 84 INeohipparion sss.4)- 2. eee 1.17

te WOMNOOAMN soon0420c 97 Equus caballusi i. <ck. 52 eee a2,

BUMPLOLOP OMIA nike ie eed ee 1.01 RS SCOttl 4. 25.5 ee a eee 88

Meniscotherium... ............ Oil Mra oulus.ei4oo cine Sere ee 1.09

ROGCAVIA a ears orm; ortsiont oe as 2 sy Odocoleus:).22.).4.¢< 3.5 42 1.16

IbfiyraChy Usins cea oe at eee eae 95 Gazellatci) Weisiaa.. oe ee « 125

SUSTNCLOfate aaa ote ere Spe ee .86 Antilope. co. .455 a. - ee eee 1.21

HH OGYLOPUS tae ey cteiscaressyeierers See oe .96 Amtilocaprale sacl: = tei ee 1 28

These figures express the fact noted by Osborn, that, in general, gravi-

portal forms have relatively a short tibia and long femur, while cursorial

forms have a long tibia and short femur. It is highly probable that in the

remote ancestors of all the Placental orders, the tibia was long, perhaps

about as long as the femur. The primitive combination of long tibia and

short feet is retained in Meniscothertum, Hyraxv and to a less degree in

Euprotogoma, Phenacodus wortmam and Hohnppus. 'The steps through

which the shortening of the tibia in graviportal animals has been

attained may in some cases be discerned. Thus in the Amblypoda, the

small and primitive Pantolambda has a tibio-femoral ratio of .76; in

Coryphodon, this ratio falls to .61 and in the huge Uintatherwum to .53;

similarly in the titanotheres, the ratio drops from .77 in Palwosyops to ~

-54 in the massive Brontotherivwm. In the Rhinoceroses, it falls from

.95 (Hyrachyus) to .79 (Rhinoceros) and .58 (Metamynodon).

In the extremely cursorial lines, the long tibia of primitive mammals

was usually lengthened slightly, but in the heavier members of cursorial

phyla (e. g., Hquus, Bison), we observe occasionally a falling off in the

ratio.

The difference between the shoftest graviportal tibia (Uintatherium,

ratio .53) and the longest cursorial tibia (Gazella, 1.25) is much less

extreme than in the metatarso-femoral ratios, which range from .11 to

GREGORY, QUADRUPEDAL LOCOMOTION 289

1.35. There are also more ratios of intermediate type. Consequently,

the tibio-femoral ratios taken alone do not always furnish a sure indica-

tion of the mode of locomotion.

From the viewpoint of adaptation, there are several plausible reasons

why the tibia has not shortened to so extreme a degree as has the middle

metatarsal. A short tibia implies a low knee joint and a long femur.

A long femur, as stated below, is associated with a nearly vertical in-

biceps 1.’

Wic. 6.—Relations of certain extensor muscles to the pelvis and femur in the standing

pose in (I) a typically cursorial form, the Horse, with relatively wide angles

of insertion (a, B, y), and in (II) a typically graviportal form, the Mastodon,

with narrow angles of insertion (a’, B’, 7’).

The heavy black lines represent the general directions of the muscles; the broken lines

represent the radii of rotation of the insertion points.

nominate bone and with narrow angles of insertion of the principal long

muscles (Fig. 6). The result of these small angles of insertion is

that the long muscles exert a powerful pull in the direction of the shaft

of the femur (p. 278 and Fig. 2), an arrangement favorable to the

lifting and support of great weight (p.-290). The short tibia and

290 ANNALS NEW YORK ACADEMY OF SCIENCES

short foot, conditioning a low knee joint, also enable the very heavy

animal to rise from the ground with comparative ease. At the same

time, excessive shortening of the tibia (as in Ground Sloths) probably

conditions very slow motion, which would be very disadvantageous to an

animal that has to wander far in search of food.

GRAVIPORTAL AND CURSORIAL TYPES OF FEMUR

The femur of primitive animals (Creodonts, Condylarths, Stem Peris-

sodactyls, etc.) is long, when compared with the size of the body. It

has a prominent third trochanter situated in the upper fourth of the

shaft and serving for the insertion of the gluteus superficials (seu

maximus). This muscle arises from the enlarged tuber coxe.of the

ilium and apparently functions as an adductor of the femur. The

trochanter major projects high above the shaft.

These characters persist in the cursorial types, the chief difference

being that here, as a rule, the femur is relatively short when compared

with the size of the body. The short femur is associated with a sub-

horizontal innominate bone and with comparatively open angles of in-

sertion of the long muscles (Fig. 6, I), which pull across the long axis

of the femur and give high components of rotation and relatively low

centripetal components (in the direction of the shaft of the bone).

Such an arrangement, associated as it is with cursorial adaptations,

seems less adapted for the support of a great dead weight than that no-

ticed above in graviportal animals. In the standing pose, a sharp angu-

lation at the knee conditions a tendency for the leg to collapse, which is

counteracted by the stretching and tension of the opposing flexors and

extensors of the thigh (Fig. 7). In the horse, these opposite tensions

are transmitted by means of special tendons on opposite sides of the leg,

which also serve to tie as it were the limb in position and thus relieve

the muscles to a considerable extent. In graviportal animals, the long

muscles are also stretched in standing and thus serve as ligaments, and

since they are inserted at very narrow angles, their centripetal compo-

nents are relatively high (p. 278).

In the femur of graviportal animals, the third trochanter is often

reduced, absent or confluent with a long ridge running down from the —

great trochanter and situated well down the shaft. The functional

meaning of this is possibly that the gluteus superficialis is less developed

in graviportal animals, its place, perhaps, being usurped by the greatly

enlarged gluteus medius. Again, it is conceivable that the third tro-

chanter may have been crowded out, so to speak, by the enlargement of

the powerful vastus muscles, which arise on the femur just above the

GREGORY, QUADRUPEDAL LOCOMOTION 991

third trochanter and which are one of the chief muscles used in lifting

the body at the hips. The great trochanter of graviportal animals is

very broad and heavy, but is more or less sessile, not jutting up so much

tensor

fase 1ae

tendo accessoriws

oa from biceps and

semitendinosus

Resultant pull, tendons

of knee and ankle:

a) Flexion: tendo femoro-

tendon, biceps. Vso uss

to metaca rpus

i Z

¥ie. 7.—fhe tendons and muscle-pulls which hold in place the fore- (1) and hind- (11)

limbs of the standing horse

After Schmalz

above the head of the bone. The reason is that the massive gluteus

medius is inserted more upon the flat top and outer side of this projec-

tion than upon its anterointernal slope (Fig. 6, II), a fact correlated

with the subvertical position of the ilium.

299 ANNALS NEW YORK ACADEMY OF SCIENCES

In brief, the long femur of graviportal animals serves for the attach-

ment of long and very massive flexor and extensive muscles inserted at

slight angles; it brings the knee joint well down below the belly, and it

is further characterized by a very heavy sessile great trochanter and a

more or less reduced third trochanter, located far down the shaft; the

distal facets, as observed by Osborn, are more or less at right angles to

the shaft, so that the femur rests subvertically upon the tibia.

GRAVIPORTAL AND CURSORIAL TYPES OF PELVIS

(Figs. 4-6)

The upper anterior border of the ilium in cursorial animals is con-

cave, so that the anterior part of the gluteus medius is produced in front

of the ilium and inserted into a long fossa in the posterior end of the

fascia covering the longissimus dorsi.*? The result is that these two

muscles apparently pull in tandem, while the subhorizontal position of

the pelvis permits the gluteus medius to be inserted on the great

trochanter at an angle approaching a right angle. Thus the powerful

and prolonged contraction of the combined gluteus medius and longis-

simus dorsi is exerted well across the long axis of the femur, producing

a very powerful rotation component. Similarly the subhorizontal posi-

tion of the ischium opens out the angles of insertion of the three

branches of the biceps and of other extensors and flexors of the thigh.

In such progressively graviportal lines as the Amynodontine among

the Rhinoceroses, we observe the transformation of the cursorial into

the graviportal pelvis by the filling out of the anterosuperior concavity

of the ilium, the broadening of the ilium and outward growth of the

tuber coxeze and the increasing verticality of the whole pelvis. The

result of these changes with regard to the muscles is that the gluteus

medius is separated by a wall of bone from the longissimus dorsi, the in-

sertion space of the gluteus medius and accessorius is greatly increased

and the medius passes down subvertically on to the broad trochanter

major of the femur; the angles of insertion of the flexors and extensors

also lessen. Such graviportal adaptations have taken) place independ-

ently in many lines of ungulates (Amblypoda, Proboscidea, Titano-

theres, Amynodonts, Toxodonts) and also in the ground sloths.

422R. ScumMautTz: “Atlas der Anatomie des Pferdes.” Zweiter Teil : Topographische My-

ologie, Taf. 27 u. 28. Ato, Berlin, 1909.

GREGORY, QUADRUPEDAL LOCOMOTION 993

GRAVIPORTAL AND CURSORIAL TYPES OF FORE LIMB

Much of what has been said above regarding the mechanical advan-

tages of short hind feet in moving a heavy load or of long feet in throw-

ing forward a light load applies also to the fore feet. The table of ratios

(Plate XXXIV), however, shows that in both graviportal and cursorial

types, the third metacarpal is usually longer in proportion to the radius

- than is the third metatarsal to the femur. The radius also is usually

much longer in proportion to the humerus than is the tibia to the femur

and the radiohumeral ratio responds less clearly to changes in mode of

locomotion ‘than does the tibiofemoral ratio. The adaptive reasons for

these facts are not altogether clear, but they must be conditioned partly

upon the marked differences in form and musculature between the shoul-

der girdle and the pelvis and upon the different situations of the fore

and hind limbs with respect to the center of gravity of the animal. The

relatively long metacarpals and long radius even of graviportal animals

gives a very long reach to the fore arm, as shown in Fig. 1, I. In

some graviportal animals, as in the Proboscidea, the humerus is much

longer than the scapula, the elbow is widely exserted from the body and

is turned outward in the back stroke, thus bringing the wrists well in.

Interference is avoided by the sharp pronation of the radius, so that

during flexion of the wrist the palm of the manus is turned partly out-

-ward. In this type of scapula, the backward extension of the posterior

_angle of the blade, as well as the length of the humerus, gives space for

a very heavy caput longus of the triceps, which thus also secures a more

direct upward pull on the olecranon. The postspinous fossa is located

almost directly above the posterior part of the great tuberosity of the

humerus, so that the pull of the infraspinatus muscle is nearly vertical.

' The tuber spine of the scapula is greatly enlarged, its upper border for

the trapezius, its lower for the heavy deltoideus. In some other gravi-

portal types (e. g., Tovodon, Rhinoceros), the scapula instead of being

broad and short is narrow and high, with vertically extended pre- and

postspinous fosse. In some cursorial types (e. g., Hquus), the scapula

is also long, with rather narrow pre- and postspinous fosse; in others

(e. g., Pecora), the scapula is fan-shaped, truncate at top, with pre-

spinous fossa reduced and postspinous fossa much developed.

Regarding the relative power of the fore and hind limbs, many people

think that in the horse, the hind limbs furnish much more than half of

the locomotive power, but Stillman arrays cogent evidence** tending to

#£ Op. cit., pp. 69, 79, 89.

994. ANNALS NEW YORK ACADEMY OF SCIENCES

show that the fore feet, drawn backward by the great pectoral triceps

and powerful back muscles, contribute more than their share to the

general result.

DIAGNOSTIC VS. CONVERGENT EVOLUTION VALUE OF LIMB RATIOS

From inspection of the table (Plate XXXIV), it will be seen that,

allowing for imperfection of the fossil material and incompleteness of the

series, the limb ratios have a certain degree of diagnostic value when

taken in groups and that pure convergent evolution rarely brings about

a close agreement in all four ratios at once.

The best cases of convergent evolution noted are as follows:

Mts. IT T. . “Mite ieee

F. Ds, H H

Girseal Perissodactyla ...Neohipparion...... 1.01 TaN 1.16 1.30

*°* | Artiodactyla..... Odocoileus......... 1.00 1.16 1.05 1.12

: Hdentatar 2.2. Lestodon 12 Ol 17 .60

Rraviportal. { Amblypoda..... Coryphodon....... 14 61 5S) 66

{ Titanotheriide...Brontops robustus.. .26 00 .o7 .82

Te | Amynodontide..Metamynodon..... 24 .58 .o9 .81

Mediportal. Rhinocerotide. ..Teleoceras.......... . 25 .O7 37 78

Toxodontia...... Moxodomye ee nelays elZ .06 .38 ahi

The greatest extremes are found in the following: —

Traviportal. Amblypoda...... Uintatherium...... 10 .o3 .19 .70

Cursorial... Giraffide........ Giralttay iia. cases 1.35 1.18 1.42 1.60

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Annas N. ¥. ACAD, Sct. Vonume XX, Prats XXXIV

COMPARATIVE TABLE OF LIMB RATIOS IN THE HOOFED MAMMAIS.

= ns = a=! rt

= aS i > aq = Jl ox =

& an ~ = i 2 an = a Fr

Species. = ta =| ah P z a3 = @ I

x 82 =) = & o ss = 5 |

= ss al & aot g ap ia 3 o

5 oS 2 a) | 5 oC 2 =

ene 2 a | oe) a | 8 = a | 3

CoNDYLARTHRA MM. mum. mm, mm mm atl

Euprotogonia puercensis ....-...... -| 105 45 43 107 ; er SLEDS Os enorCor esonccas|! Sakurce|| accocn

Phenacodus wortmani...........-..-. 134 51 .38 { ne r| 1.00 \ 107 36 65 89 83

Phenacodus primzevus ...........+--- 234 74 31 198 -84 167 70 42 146 .87

Meniscotherium terreerubrie........-- 100 2 -29 91 ht 82 22 .27 58 .70

AMBLYPODA

Pantolambda bathmodon............. 149 36 .24 114 -76 124 31 .25 82 .66

Coryphodon lobatus ........... ....- 423 62 14 260 -61 363 70 19 240 .66

Uintatherium (Dinoceras) mirabile...| 692 70 -10 360 53 540 106 at) 380 -70

PYROTHERIA

Pyrotherium (figd. by Gaudry).......| 622 | ......]...... 351 -56 Cb | ceAnina [oso vor 238 52

PROBOSCIDEA

Mastodon americanus................ 1,020 e 117 Al 705 69 885 165 18 670 .75

Boab ites bth eS arasarnaccues OC 1,020 138 13 618 .60 810 183 -22 685 -80

KE. (Loxodonta) africanus............ 1,050 144 13 755 7i | 1,000 205 20 870 87

HyRacorDEA

PSOE ANE Oran lita lotatetercSalst= sinlvisirieiaialeiatst 71 19 .26 69 Ne/ 69 16 -23 46 .66

ToxODONTIA

ALOR OGOURADemerietateislcisthicter feleinicnterarereratele 577 101 Abt 325 56 387 147 .33 298 77

EDENTATA GRAVIGRADA

lab ferilo)s):01:\ os see once dpe OOnDueOUmE ae 150 18 12 108 :72 13 22 16 106 .80

estodon armatus,......2........-0.06 640 78 12 330 51 530 94 17 325 -60

PeRISssODACTYLA TAPIROIDEA

Heptodon calciculus ............... at) ude 75 e 43 175 1.00 115 67 -58 114 -99

Tapirus americanus - | 262 108 -41 208 ‘79 205 106 -50 177 .86

AA OMIE Eh UNC A gooadconoanounonor 320 120 -37 258 -80 250 120 48 228 91

PrrissopacryLa RaiNoo:

Hyrachyus agrarius.... 254 110 43 243 95 197 93 47 197 1.00

Hyracodon nebrascensis c 267 114 42 220 -82 202 114 56 210 1.03

Rhinoceros indicus...............+-+ 495 180 .37 395 -79 385 186 48 385 1.00

Teleoceras fossiger........0:..sce00 408 105 .25 233 57 305 e 114 -37 238 -78

Metamynodon planifrons............- 480 118 :24 280 .58 393 103 39 320 81

PerissopAcTYLA TITANOTHEROIDEA

Hotitanops borealis .........-........ PANG |) eae easub |p enuees 203 tf} a ieee 1 eal erie ns A [oer cr ene

Paleeosyops major..............2sse0es 433 137 31 832 ED revste eayeel | petmeetoees teal (catetarereta || eaarertene

Palseosyops leidyi.........--...--..-. 370 110 .30 290 -78 325 2113 235

IFAP Nose ose Ae loan ola blnkep PsP eigen ll asnediellesetine \jesrnnoaa lPoaceen 2340 106 237

Limnohyops sp. (A. M. N. H. No. 11689} 35 e 111 31 285 -79 293 109 228

ae st fs ** No. 11690 387 123 31 283 BA: Se eeavesinel becca: |jeonece Crd sme. Gl

Manteoceras manteoceras............. MGR Vitsantas secon s 272 Bok Wes eee Aen ese mcicatie eocsier. on feat os e

Mesatirhinus petersoni, No. 11659. 308 118 33 283 A Mela S Seis apeane.||RRaonode i neemaa

Dolichorhinus hyognathus, No. 13164.| 386 119e OO eS fctevateie) || Beyaratese LONE) |Meat 284

Titanotherium trigonoceros........... 770 220 e -28 430 55 620 240 520

Brontops/robustus?.. ses. 010s emesis 812 212e .26 448 As 608 230 504

Brontotherium gigas 9 (518 A. M.)...] 780 200 -20 427 54 528 214 78

PERISSODACTYLA HiPPOIDEA

OHIDPUSIAD saeraneien sees eck cser 162 82e -90 162 1.00 121 64 53 110 -90

MGR OpodiY OWI eh moagascangsucHesonuod 178 121 .68 193 1.08 136 92 .68 136e} 1.00

Hypohippus osborni.................. 278 218 -78 277 1.00 205 203 -99 260 1.27

Neohipparion whitneyi ... ats 249 252 1.01 293 1.17 187 218 1.16 244 1.30

IQUE KAN Over a eee tore area 313 277 .88 310 +99 237 238 1.60 302 1.27

UGUIMISISCOLEL eye sternisrmtaicterabs enisrereds sterner 370 263 71 330 .88 289 243 84 342 1.18

Equus caballus (race horse).........-. 392 288 -73 363 92 305 240 -78 363 1.19

Hippidion neogeeum................- 340 214 -62 305 .89 273 198 :72 287 1.05

ARTIODACTYLA

Oreodontidxe

= aoredon Cul bextSOnie sents 161 62 .38 142 .88 138 57 41 113 81

wide

SUSI RCROL ese rces cre viele Sistelere Aes 248 86 -34 216 .86 208 77 .37 168 -80

Hippopotamids

Hippotamus amphibius.......... 498 130 -26 332 -67 395 152 -38 270 .68

Camelidee

Kotylopus reedi.................. 148 78 52 142 :96 118 68 57 100 .84

Camelus arabicus................ 470 325 .60 400 -80 363 330 -90 455 1.25

Tragulidx

AUEMUE GAN eS docseonussaaanane 94 62 -66 103 1.09 74 42 -56 62 83

Cervidee

Odocoileus hemionus............. 253 255 1.00 295 1.16 198 208 1.05 223 1.12

i Genie MMOPACBLOS <1 crane ereicinter cine 430 350 -71 454 1.05 334 342 1.02 358 1.07

ovidse

Gazella dorcas juv. .............. 140 132 81 176 1.25 93 134 1.44 118 1.26

Antilope cervicapra.............. 183 183 1.00 223 1.21 133 180 1.35 168 1.26

IBINON) DISON. incisive oe eye aveutere 369 243 -65 355 -96 290 198 -60 293 1.01

Antilocapride

Antilocapra americana........... 210 218 1.03 260 1.23 164 213 1.30 202 1.23

Giraffide

CPE CT hls ee geScorraneeece 2.) 466 630 1.35 550 1.18 4385 618 1.42 698 1.60

e—Estimated.

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| Editor, EpMUND Oris Hovey

AGE OF THE BEDFORD SHALE

OF OHIO

Grorce H. Girry

Bo EW YORK

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_[Anwnats N. Y. Acap. Sciences, Vol. XXII, pp. 295-319. 13 November, 1912]

GEOLOGIC AGE OF THE BEDFORD SHALE OF OHIO?

By Grorce H. Girty

(Presented in abstract before the Academy, 7 October, 1912)

The Bedford shale takes its name from Bedford, in Cuyahoga County,

Ohio. The typical locality is the gorge of Tinkers Creek below the falls,

where the formation comprises about 75 feet of bluish clay shale lying

between the Berea sandstone above and the black slaty Cleveland shale

below. To the west, the lower portion of the Bedford develops a sand-

stone member which is quarried as the “Cleveland bluestone,” while the

upper portion undergoes a change of color to a strong red. In fact, the

Bedford is better known as a red than a blue formation.

The fossils of the Bedford shale are largely confined to the basal por-

tion, though a few species are represented by rare individuals at higher

horizons. At Bedford, fossils are abundant in immediate contact with

the Cleveland shale, where they are more or less crushed, and also a few

feet higher in large calcareous concretions, where the surface characters

are apt to be obscured. Some of them are broken and rounded as if by

wave or current action, but this is not the general character of their

occurrence.

The Bedford shale lies close to what has been considered the boundary

between the Devonian and Carboniferous systems, and it is the purpose

of the present paper to present such evidence as I have bearing on the

geologic age of the formation. The question then is whether the Bedford

shale shall be included in the Devonian or the Carboniferous system. I

shall treat this largely as a paleontologic question, and my fossil evi-

dence is derived from typical sections at Bedford and other points in

Cuyahoga County.

Tt will not be out of place to consider some of the principles controll-

ing such an attempt as I have taken in hand.

Theoretically, the great geologic systems were defined by movements cre-

ating extraordinary changes in the conditions of land and water, always

undergoing changes more or less gradual, and these conditions entailed

corresponding changes in the character of the plants and animals which

had in them their habitat. As expressed lithologically, the rocks of the

several systems, perhaps generally in the typical region and not infre-

1 Published by permission of the Director of the U. S. Geological Survey.

(295)

296 ANNALS NEW YORK ACADEMY OF SCIENCES

quently in regions far removed, show a basal sandstone or conglomerate

with more or less pronounced evidence of preceding erosion or uplift.

Such events though often far reaching must in the broadest sense have

been only local, but we may fairly expect the more subtile changes of

environment, such as depth, temperature, currents, which govern the

character and distribution of animal life, to have extended far beyond the

area of great disturbance, and we may look for evidence of such disturb-

ance in the fossil floras and faunas, when none is to be found in the rocks

themselves or their relation to one another. In fact, such paleontologic

evidence may have been all but universal. Conglomerates and uncon-

formities and faunal changes occur at other horizons than the division

lines between systems, but such evidence will be of great importance in

deciding the point in question.

Yn addition to the kinds of evidence already mentioned as often accom-

panying the transition from one geologic system to another, that is, an

interval of erosion, a basal sandstone or conglomerate and a well-marked

faunal change, there are also certain other considerations of a more ad-

ventitious or incidental nature. For practical purposes, it would be

unfortunate if this line (that between two systems), which of all lines it

is desirable to represent on a geologic map, were taken where it could

with difficulty be recognized in the field, as in the middle of a uniform

lithologic interval, or where the evidence would often be concealed, as

would be the case in some regions if it were assumed to he between two

formations of soft and easily disintegrated material. Furthermore, the

importance of convention also enters the consideration. Other things

being equal, it is clearly preferable to take for the boundary which is

sought the same horizon at which it has been drawn elsewhere, if that

horizon can be determined.

The boundary between the Berea sandstone and the Bedford shale in

some degree satisfies all these requirements. It is easily recognized and

easily traced; it also appears to be the locus of an unconformity.” The

Berea answers to the basal conglomerate of theory. While the Bedford

seems properly to form part of the great shale series which preceded it,

the Berea marks the change to another and different type of sedimenta-

tion. The passage from Bedford to Berea is also marked by an abrupt

and strong faunal change. Not only are the two faunas widely different,

but the Bedford has a preponderant Devonian facies and the Berea a pre-

ponderant Carboniferous facies. To some extent, the faunal change at

; 2 This erosional unconformity was clearly stated by Newberry as early as 1874 (Ohio

Geol. Survey, Geol., vol. 2, p. 91). More recently it has been mentioned by J. E. Hyde

(Jour. Geol., vol. 19, 1911, p. 257) and described by W. G. Burroughs (idem, p. 655).

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 997

the top of the Bedford is offset by one equally marked in the opposite

direction, between the Bedford shale and the Chagrin shale. In support

of this statement I shall not feel called upon to submit evidence, because

in so far as it is untrue, the Devonian affinities of the Bedford faunas are

by so much stronger than they are here represented. Finally, the deter-

mination of the boundary between the Devonian and Carboniferous at

the base of the Berea satisfies to some extent the canon of convention.

This is true, however, only from one point of view, for if the Berea

(“Corry”) sandstone correlates with the Kinderhook beds of the Missis-

sippi Valley which represent the base of the Carboniferous system in its

type section, and if some horizon below (?) the Chagrin correlates with

the Chemung group which represents the top of the Devonian in its type

_ section,*® the canon of convention or usage would be ambiguous in its

bearing upon these intermediate beds whether they should be classed with

one system or the other, because by one approach they would be found

above the recognized top of the Devonian, just as by the other they would

be found below the recognized base of the Carboniferous.

This too is only partly true, however, because Hall* classed as Che-

mung, or more often “Upper Chemung,” the intermediate group of

strata here under consideration, so that, although their stratigraphic

position is above the Chemung proper and therefore theoretically above

the top of the Devonian, they have in practice been included all along in

that system. Incidentally, this circumstance seems to show Hall’s opin-

ion of the affinities of the “Bradfordian” faunas,° though it must be re-

membered that he correlated with the Chemung the Waverly group lying

still above.

Now the statements which it is proposed to present the evidence for

and to discuss are the following: the faunal change incident to the pas-

sage from the Bedford shale to the Berea sandstone; the Carboniferous

aspect of the Berea fauna, and its probable correlation with the Kinder-

hook of the Mississippi Valley; the Devonian aspect of the Bedford

fauna, and the relation of the Cuyahoga, Berea and Bedford faunas to

those of the typical Mississippian sections of Missouri, Illinois and Iowa. |

%The typical sections of the Devonian and Carboniferous in the United States are

meant. The original sections are of course in Hurope.

4This statement is based upon the fact that Hall described and included in the Che-

mung fauna numerous pelecypods from many of the well-known “Bradfordian” locali-

ties of Pennsylvania.

5 Here and elsewhere in this paper, ‘‘Bradfordian’”’ is employed in the sense in which

I defined it in 1904 (Science, nN. s., vol. 19, p. 24)—-for a series of strata about 500 feet

thick, lying between the typica] Chemung and the Waverly group (as limited below by

the Berea sandstone), and comprising in the typical sections near Bradford, Pa., and

Olean, N. Y., the formations described under the names of Cattaraugus, Oswayo and

Knapp. :

998 ANNALS NEW YORK ACADEMY OF SCIENCES

Some diversity of opinion exists as to the principles which should

govern the interpretation of paleontological evidence in determining the

boundaries of geologic systems. JI propose to consider these different

views and to decide which is to be employed in the present investigation.

Thus, to take a put case, it is sometimes said that the line between the

Devonian and the Carboniferous should be placed at the first introduc-

tion of Carboniferous species. This principle seems by implication or

otherwise to be adopted by Glenn and Butts and Clarke in their discus-

sion, of the geology of the Olean quadrangle and other areas in western

New York.® It is true that, in adopting this principle in the Olean rock

section, the authors were influenced by the belief that the interval below

the Olean conglomerate member of the Pottsville formation in the Olean

quadrangle corresponded to the interval similarly underlying the Olean

conglomerate in northwestern Pennsylvania and northeastern Ohio, so

that several hundred feet in the one section would have to correspond to

the authentic Waverly group in the other.*. Now that it is known, or at

least seems highly probable, that, owing to erosion which preceded the

Olean conglomerate, that member rests on lower and lower strata as it is

followed eastward, so that at Warren, Pa., most of the Waverly rocks

are missing and they are not known to appear-in the sections farther

east,—with this condition of affairs granted, it is possible that the au-

thors would not have adopted the principle of first appearance. It is

necessary, however, to examine this principle to see whether it is gener-

ally applicable or applicable to the present case.

The meaning of the principle as stated above is clearer at first sight

than when it is examined more closely. The implication largely turns

on, the meaning which is given to the terms “Devonian” and “Carbon-

iferous” species, and, at the risk of drifting into something like the

Greek dialectic, it will not be unprofitable to consider this question. By

Carboniferous species may be meant (a) any species which has been

found in rocks of Carboniferous age; (b) a species especially abundant

or widely spread and persistent in rocks of that age though occurring

elsewhere, or (c) a species known only from the Carboniferous, of at

least known there but not in the other geologic system with which com-

parison is made.

It is evident that, in the present instance, the words “Carboniferous

6 New York State Mus. Rept., vol. 56, pt. 2, 1903, pp. 985, 991 and 999. Clarke gave

a somewhat more extended discussion the year previous (idem, vol. 55, 1902, p. 524) in

which he showed that a marked faunal change took place at the base of the “‘Brad-

fordian,”’ but not that the ‘“Bradfordian’’ fauna had a preponderating Carboniferous

aspect.

" Loc. cit., p. 991.

GIRTY, GEOLOGIC AGH OF THE BEDFORD SHALE 299

species” cannot be taken with the first meaning (a), because some types

found in the Carboniferous have a very long range and originated at

much lower horizons. Leptena rhomboidalis is an example.®

If we assume the second meaning (b) for the expression “Carbon-

iferous species,” the principle would mean frankly a redetermination of

the Devonian-Carboniferous boundary, not only in other areas but in the

typical area as well, a redetermination, moreover, which would never

cease, because the Devonian beds which would thus be added to the Car-

boniferous would carry over other species having a sporadic appearance

at lower horizons. This would entail a new adjustment and so on appar-

ently until the bottom of the stratigraphic column was reached. Fur-

thermore, is it one such species, or two, or a score that the application of

this principle involves? Reasonably but one. The decision might easily

then come to depend on the identification of one or two specimens re-

sembling several closely related species.

If we now take the last meaning (c) for the expression “Carboniferous

species,” 7. e., species characteristic of the Carboniferous, those not known

to occur at any horizon outside of the Carboniferous, the statement under

consideration becomes hardly more than a truism, but a truism which

assumes that we have complete knowledge of the range of species, that

the species in question are not only not, known to range outside of the

Carboniferous but that they cannot so range, and this is an assumption

which everyone knows is quite inadmissible, since the range of species as

recognized at any time is continually being changed by new data. Other

considerations might be brought forward, but it is already plain that this

is not a workable principle, no matter how understood, for determining

the boundaries between systems, or for classifying the Bedford shale.

It might be held, on the other hand, that the proper way to fix the

line between the Devonian and Carboniferous systems is by the disap-

pearance of the last Devonian forms. This principle is the antithesis

of that just considered. It is open to the same objections and is equally

untenable. |

The only practicable method of interpreting paleontologic data in

most cases of this sort is evidently by taking the balance of evidence. By

this method the decision hangs not upon one or two forms but upon the

entire number known, and, although the evidence of each form may be

impaired by poor material and close relationship between species, it be-

comes cumulative. Here the expressions “Carboniferous” and “Devo-

nian” species signify species especially common in their respective sys-

8 De Koninck, however, distinguishes the Mississippian Leptzena as a distinct species

under the name Leptena analoga.

300 ANNALS NEW YORK ACADEMY OF SCIENCES

tems. If the species are absolutely unknown outside of their respective

systems, the evidence is so much the stronger, but the difference is one

of degree only, and no assumption is made that a species at present known

in the Carboniferous may not subsequently be found in the Devonian or

vice versa. 'That a species has not yet been found in the other system,

however, rightly gives it exceptional weight especially in regions where

the available data on range are considerable.

This principle is of course unsatisfactory. It is impossible to estimate

or to state the results mathematically. Different species have different

values, and the same species may have different values in different re-

gions. While these values are not expressible in numerical terms and

indeed must vary somewhat with each observer according to the character

and extent of his experience, nevertheless, in practice, the evidence is

seldom so nicely balanced or the antecedent experience of those who judge

it so diverse that reasonably satisfactory and unanimous conclusions are

impossible.

In determining the relationship of geologic formations, which includes

also the determination of their geologic age, some species have, as already

pointed out, greater importance than others. This is partly because we

know more about some than others. The determination of such rela-

tionships as I have mentioned rests very largely on our knowledge of the

range in geologic time of different types of fossils and involves one of the

most fallible of all processes of inductive reasoning. Because a fossil

has not yet been found above or below a certain horizon it does not follow

that it never will be so found, yet that is virtually the inference on which

all correlations and age determinations are based. At best, this fur-

nishes conclusions which are fairly safe, and at worst it furnishes con-

clusions which are highly unsafe. At its best, the conclusion depends

upon the concurrence of a large number of species and upon species

whose range has been ascertained by a large number of observations.

For the same reason, it is clear that common species are more significant

than rarer ones, because our knowledge about their range is more trust-

worthy, and, in some cases, fairly sound inferences can be drawn from a

single species.

It is, however, not only the trustworthiness of our knowledge which

lends greater significance to some forms than to others, but also the

length of range in geologic time, which differs with almost every form;

for, obviously the presence of a form which had an established range of

100 feet would be much more significant in correlation than that of one

whose range was 1000 feet. Here enters also the consideration of groups

larger and smaller than species, the range of which is, generally speak-

GIRTY, GHOLOGIC AGE OF THE BEDFORD SHALE 301

ing, in proportion to their size,—long as the biologic rank of the eroulp

increases, short as the biologic rank of the group decreases.

Our knowledge of the range of fossils is conditioned by other peti

than the amount of data as to their occurrence, since it manifestly in-

volves the identification of the fossil as a species and the identification of

its geologic horizon. Another factor then which makes some forms

more significant than others is that their generic and specific relations

can be determined with greater certainty. Variation in this particular

is in some cases intrinsic, in others extrinsic; often it depends upon

both factors. Manifestly, in groups where variation is restricted in

degree, or is restricted to a few characters, or is marked by complete

intergradation, discrimination of species and even of genera is more

difficult than others. ‘Types which vary in shape alone have in my ex-

perience proved especially unsatisfactory, while those which are sculp-

tured or which possess other features of relief give more reliable results,

especially when, as is almost always the case, this is combined with varia-

tion in configuration also. Preservation, which in fossils has always

destroyed the soft parts and the coloration, often obscures other charac-

ters too, and this deterioration, owing to peculiarities inherent in whole

groups of shells, is more liable to befall some types than others. As is

well known, owing to their physical rather than their chemical structure,

the shells of pelecypods and cephalopods are apt to be removed by solu-

tion so that in Paleozoic rocks they are as a rule reduced to the condi-

tion of molds, while the shells of brachiopods retain their original compo-

sition. Also, partly because of the solution of the shell, which tends to

obscure both sculptural or specific and structural or generic characters,

partly because of being marked by growth lines alone and having the

generic characters largely developed along the hinge where they are at

best difficult of observation, many types of pelecypods show differences

only in shape and configuration (which are peculiarly liable to be altered

by compression), while at the same time quite widely different genera

are in general aspect, which is about all that can be determined, very

similar,—for these reasons pelecypods often prove an unsatisfactory

eroup for stratigraphic paleontology, since even the generic position of

specimens as they ordinarily occur in the Paleozoic is often undeter-

minable except on characters which are not of themselves strictly generic,

although they are, or appear to be, correlated with generic characters.

It is obvious that, while the larger zodlogic groups are as a rule of less

value because of their longer range, they are at the same time usually

of greater value because of the precision and certainty with which they

can be distinguished, for it hardly needs to be stated that a specimen

302 ANNALS NEW YORK ACADEMY OF SCIENCES

can often be referred to a genus with certainty while its specific position

is a matter of doubt. On the other hand, a provisional specific identifi-

cation is sometimes only possible on the assumption of a generic one.

Though because of their more extended range (in part compensated by

the certainty of delimitation), the larger groups are less serviceable in

correlating different sections, they are more instead of less valuable in

estimating the importance of faunal changes in the same section, since

they indicate a greater degree of change and possess the added advantage

of increased certainty of discrimination.

Of equal importance with identification of a form in its biologic rela-

tions is the identification of its geologic horizon and this is frequently

unsatisfactory. It seems to be true, and it is natural that it should be

so, that in geologic time, as at the present day, the progress of sedimenta-

tion and the course of biologic development varied in different areas or

provinces, and that deposits may be, so far as one can tell, essentially

contemporaneous, and yet very different in lithologic character and in

the character of their fauna and flora. The limits of geologic provinces

are not clearly defined, if indeed it is not ultimately shown that they are

without definite limits but are continuous with one another; and still

less is it known what were the factors which produced their differentia-

tion. For my own part, I have very little faith in the theory of barriers

(in the sense of land barriers) as a panacea for all the ills of strati-

graphic geology. On the contrary, I believe that during geologic time,

as today, the conditions controlling the character and distribution of

faunas are depth, temperature, food supply, current action, salinity, bot-

tom and so forth. At all events, as between different provinces, most

correlations are at present more or less provisional, so that while the

paleontologist must not disregard the data from any area, his deductions

concerning one province should be largely guided by the data from that

province.

Since the data of range and distribution of species are in large meas-

ure not on record even when they have been ascertained, and since the

records are very scattered, each investigator must approach a problem

with a different store of facts on which to base his inference as to geo-

logic age and correlation, but it by no means need follow that such partial

or even one-sided knowledge must lead in different cases to different

conclusions.

A number of years ago when much engaged with the investigation of

these “Bradfordian” beds, from which I have since been temporarily

diverted, I began and all but completed a descriptive study of the fauna

of the “Corry” sandstone (since correlated with the Berea sandstone).

GIRTY, GHOLOGIC AGE OF THE BEDFORD SHALE

‘This fauna I will now list in the terms in which it was then prepared

as follows:

Crania levis Keyes

Rhipidomella n. sp.

Schuchertella desiderata Hall

Clarke

Producitella n. sp.

Productus vn. sp.

idem, n. var.

n. sp.

arcuatus Hall ?

levicosta n. var.

Strophalosia-like form, n. gen. n. sp.

Spirifer marionensis Shumard

disjunctus Sowerby ?

Cyrtina triplicata Simpson

Syringothyris angulata Simpson

extenuata Hall

and

303

Camarotechia metallica White ?

Paraphorhynchus striatum Simpson

mediale Simpson

Pterinopecten alternatus Simpson

Aviculipecten equalatus Simpson ?

patulus Hall ?

cancellatus Hall ?

Paleoneilo sp.

Sphenotus sp.

Sanguinolites senilis Herrick

Spathella ? sp.

Cypricardinia sp.

Mytilarea sp.

Edmondia ? sp.

Straparollus roberti White ?

Platyceras varians Simpson

dorsale Simpson

Athyris lamellosa L’tveillée

Cliothyridina squamosa n. var.

Camarotechia heteropsis Winchell

Tropidodiscus crytolites Hall

Conularia byblis White ?

If this list is compared with the one which I shall give farther on, it

will be seen how very differeht the “Corry” fauna is from the fauna of

the Bedford shale. Thus the statement that a pronounced faunal change

marks the transition from Bedford to Berea time seems amply justified.

‘The second point which I wish to make in this connection is that, for the

first time in this region, in the ascending series, we have a fauna of dis-

tinctly Carboniferous type. The “Corry” fauna contains much that is

new, but the development of species of the Productus rather than the

Productella group (though on this I do not lay much stress because of

the difficulty of adequately determining one group from the other), and

especially of a Productus of the cora type, an abundant Spirifer of the

marionensis type, Athyris lamellosa, a species of Cliothyridina, two spe-

cies of the Kinderhook genus Paraphorhynchus and a few other forms,

identify this horizon as Carboniferous and probably Kinderhook. There

is, to be sure, some evidence pointing the other way, as for instance a

Spirifer doubtfully identified with S. disjunctus and the two Aviculipec-

tens, also doubtfully referred to Devonian species, but no one will ques-

tion on which side the evidence is stronger.

The fauna of the Bedford shale has never been described in full. In-

complete faunal lists have been given in two or three instances. A few

species identified or figured from this formation may be found scattered

304 ANNALS NEW YORK ACADEMY OF SCIENCES

among Hall’s paleontological monographs (as Macrodon hamiltonie)

and an occasional species has been described by other writers (as Paleo-

nelo bedfordensis by Meek). MHerrick® has published a plate of figures

drawn from specimens obtained at Central College in the central part of

the State, and Foerste’® has described and figured a very limited devel-

opment of the fauna as represented in eastern Kentucky. It may be

said, however, that the Bedford fauna is very imperfectly known.

In my work on the “Bradfordian,” the fauna of the Bedford shale was

collected and in part described. As represented in my collections, the

Bedford fauna in the typical localities of Cuyahoga County, Ohio, com-

prises about 50 species, which may be listed as follows:

Lingula n. sp. Pterinopecten ?n. sp.

irvinensis Foerste ? Macrodon hamiltonie Hall

Lingulidiscina n. sp. Edmondia aff. subovata Hall and el-

newberryi Hall ? lipsis Hall

Pholidops nu. sp. Cypricardella aff. gregaria Hall and

Schuchertella herricki Foerste tenuistriata Hall

Chonetes n. sp. Sphenotus aff. cuneatus Conrad and

Productella pyxidata n. var. contractus Hall

Strophalosia sp. Pholadella newberryi Hall ?

Rhipidomella n. sp. sp.

Cranena ? aff. subelliptica Hall and Ptychodesma ? sp.

Clarke Bellerophon aff. pelops Hall, mera

Cryptonella ? sp. Hall and jeffersonensis Weller

Camarotechia sappho Hall Tropidodiscus aff. acutilira Hall, bre-

Delthyris n. sp., aff. sculptilis Hall vilineatus Conrad and cyrtolites

and mvissowriensis Weller Hall

Spirifer aff. marionensis Shumard ? Pleuwrotomaria aff. sulcimarginata

Syringothyris carteri Hall Conrad

Nucleospira ? sp. Platyceras sp.

Camarospira ? sp. Loxonema ? sp.

Athyris aff. hannibalensis Swallow Conularia aff. newberryi Winchell

and fultonensis Swallow Hyolithes sp.

Paleoneilo bedfordensis Meek. Orthoceras sp.

Leda diversa Hall Goniatites sp.

Solenopsis ? sp. Proetus ? sp.

In addition, there is a doubtful species of Rhombopora, several species

of conodonts, which are rare, and abundant though ill-preserved ostracods

suggesting the genera Primitia, Cytherella, Beyrichia (2 species) and

Paraparchites.

The identifications and comparisons given above are subject to re-

vision, but, in spite of such possible changes, the list will serve to show

® Sci. Lab. Denison Univ., Bull., vol. 4, pl. IX.

10 Ohio Nat., vol. 9, p. 515 et seq: 1909.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 205

the general character of the typical Bedford fauna as represented in very

complete collections.

Before commenting on the characters of this fauna, as shown by my

list, it will be desirable to consider some species which have been recorded

from the Bedford and which I have not identified there.

Newberry has cited the following species from the Bedford shale:

Syringothyris typa Win. ; Hemipronites crenistria Phil.

Orthis michelini Lev. Chonetes logani Hall

Spiriferina solidirostris White Lingula cuyahoga Hall

Macrodon hamiltonie Hall Rhynchonelia sagerana Win.

upon which he comments in these words:

“In this list there are several which have peculiar interest and significance,

Syringothyris typa and NSpiriferina solidirostris, for example, from the fact

that they are characteristic of the Lower Carboniferous rocks of other States,

while Orthis michelini is common to the Carboniferous formation all over our

country and in Europe.”

Herrick,’ referring I doubt not to this passage, says:

“Dr. Newberry has decided that the Bedford shale is Carboniferous on the

basis of such fossils as Syringothyris typa, Hemipronites crenistria, Chonetes

logani, Orthis michelina and Spiriferina solidirostris and a few more. Having

searched in the same localities without finding these forms in the typical Bed-

ford as it appears in southern Ohio and on the other hand finding the species

above mentioned [in a preceding list] we feel some hesitation as to the occa-

sion of the confusion. These species may indeed occur below the Berea, but

in flags and greyish shales not in the blue or red Bedford shale!”

As to the closing remark I may say that though Newberry did not de-

scribe or figure the species which he named, there is, owing to the consti-

tution of the Bedford fauna, no reasonable doubt as to what types he

wished to indicate in each case, and Herrick is quite in error in supposing

that these species did not come from the true Bedford shale.

Now, as to the species mentioned by Herrick, Orthis michelina and

Hemipronites crenistria are the species which I have listed as Schucher-

tella herricki and Rhipidomella n. sp. The difficulty of discriminating

species among the Rhipidomellas and Schuchertellas is such that these

types are of minor importance in correlating faunas. The Waverly

Schuchertellas are so closely allied to S. chemungensis that it would de-

mand considerable temerity to say that a given suite of fossils belonged

to a species of the one fauna rather than to a species of the other and

indicated either Carboniferous or Devonian age. Much the same is true

11 Op. cit. p. 109.

306 ANNALS NEW YORK ACADEMY OF SCIENCES

of the Rhipidomellas, but if reliance may be placed on the size and shape

of the muscle scars, which are usually regarded as good specific characters

in this group, I may say positively that the Bedford form is not Rhipi-

domella burlingtonensis (which was described as a variety of michelini

and is the most probable species indicated by that name which was orig-

inally applied to a European form). Chonetes logani is the form which

I called Chonetes n. sp., and which was beyond question wrongly identi-

fied with Norwood and Pratten’s species. For Syringothyris typa, I

have adopted Schuchert’s identification, S. carteri. When Newberry

wrote, and when Herrick wrote for that matter, the genus Syringothyris

was regarded as a diagnostic Carboniferous type and very justly, so far

as the facts were then known, but it has subsequently been found that

the genus occurs abundantly in direct association with Spirifer disjune-

tus in the “Bradfordian” rocks of northwestern Pennsylvania. Since

S. disjunctus has always been regarded as being as emphatic a marker of

the Devonian as Syringothyris was of the Carboniferous, it is clear that

the evidence of either type is disqualified for deciding the question at

issue. Schuchert?? has even described a species of Syringothyris from

the middle Devonian of Missouri, and furthermore a tendency to develop

the syrinx seems to be manifested in several Devonian species of Spirtfer,

so that it would seem as if the evidence of S. disjunctus should be es-

teemed of greater weight in favor of the Devonian than that of Syringo-

thyris in favor of the Carboniferous age of the “Bradfordian” strata.

Newberry’s Spiriferina solidirostris is the Deltthyris n. sp. of my list.

It is absolutely certain that this form is not the Kinderhook species

S. solidirostris and almost equally certain that it is not a Spiriferina at

all. It has, it is true, the general expression and the median septum

which are found in Spiriferina and which are also found in the group of

Spirifers to which the title Delthyris has been applied, but it does not

possess the punctate shell structure which is an indispensable character

of Spiriferina. The form in question is not rare in the Bedford shale

and I have been able to examine a considerable number of specimens.

This I have done both with a hand lens and with a compound microscope

without success in finding the punctate structure which is usually a feat-

ure easily detected in species really belonging to the genus. Thus, I am

forced to conclude that the form is not a Spiriferina, which is a typical

Carboniferous genus, but that it is a Delthyris, which is an almost equally

typical Devonian one.

I have thus traversed all the forms thought by Newberry or by Her-

rick to indicate a Carboniferous age for the Bedford fauna, and their

142Am, Jour. Sci., vol. 30, p. 223. 1910.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 307

supposed significance has for one reason or another quite disappeared

under impartial criticism.

Consideration seems to be demanded at this point of a small list of

“Carboniferous” species cited by Mr. Butts from the “Bradfordian” of

the Olean quadrangle. These species are not known in the Bedford

shale, but the Bedford interval is probably represented in the Olean sec-

tion, though not distinctly recognizable there. At all events, if the

Knapp and Oswayo formations of the Olean section are Carboniferous,

it is clear that the Bedford shale must be Carboniferous, whatever its

fauna, since if it does not represent some horizon in those formations, it

must represent one above, rather than below them. It is therefore ger-

mane to this discussion to scrutinize the evidence for calling the Knapp

and Oswayo formations Carboniferous. I suspect that the authors of the

work in which these species are cited would have proceeded differently if

they had not assumed as a postulate the general equivalence of the Olean

rock section with that of northwestern Pennsylvania and northeastern

Ohio, so that the question which they considered was not, “Is the Car-

boniferous actually represented in the Olean section?” but, “Since the

Carboniferous is represented in the Olean section, where should the line

best be drawn between it and the Devonian ?”'*

The “Carboniferous” types cited by Mr. Butts make up a total of but

seven out of a list of 59 species. All the rest are Devonian forms, most of

which, and possibly all, have never been found in rocks of Carboniferous

age, so that were we to consider the question whether the faunas show a

predominating Devonian or Carboniferous facies, there could be but one

answer. It is only by adhering to the rule of “first appearance” that

these formations can with any justification be called Carboniferous.

Let us, however, consider the Carboniferous character of the seven spe-

cies on which this age determination depends. In addition to two fishes

referred to the Carboniferous genera Ctenodus and Gyracanthus, the list

includes five invertebrates. These are Oehlertella pleurites, Orthothetes

crenistria, Glossites (Sanguinolites) amygdalinus ?, Sphenotus eolus ?,

Crenipecten winchelli.

I have already expressed the opinion that but little reliance can be

placed upon the Schuchertellas in matters of correlation because of the

difficulties of drawing any satisfactory lines between species or supposed

species in the genus. The significance in the present instance is still

13This reference of the Knapp and Oswayo formations to the Carboniferous has re-

cently been reaffirmed by Hartnagel (New York State Museum, Handbook 19, p. 87

et seq., April 1912), without the discussion that would seem to be demanded by the

subsequently known fact that these formations occur below the Waverly group of Ohio

(at least if the Berea sandstone is taken as the base).

308 ANNALS NEW YORK ACADEMY OF SCIENCES

farther reduced by the fact that Schuchertella (Orthothetes) crenistria

is a strictly European species, though the name has been applied to sev-

eral forms in the Waverly which might be regarded as a single species or

split up on rather small differences into several, according to the dispo-

sition of the investigator, and of which some are doubtfully distinct even

on trivial characters from the common Chemung form Schuchertella che-

mungensis.

From my own experience if any brachiopods are less satisfactory for

identification and therefore for correlation than the Schuchertellas, they

are the Discinoids, to which Oehlertella pleurites belongs. On the whole,

however, this species, which is a rather common “Bradfordian” type,

must be regarded as Carboniferous rather than Devonian in its bearing.

The Pectinoids are greatly diversified in shape and especially in exter-

nal ornament, and a number of fairly distinct groups can be made on

superficial characters, some groups small and peculiar, others more com-

man and generalized. To the commonest and most general of these,

Crenipecten winchelli belongs. Now species superficially very similar to

this are found in other genera, such as Pecten, Aviculipecten, Deltopec-

ten, so that, as the hinge characters are very seldom to be observed, it is

usually impossible to determine with certainty the generic group to which

these commonplace Pectens belong, and this fact naturally brings into

doubt the specific identification even when the superficial resemblance is

close. The identification in this case is made without a query, however,

and Crenipecten winchelli must be regarded as a distinctly Carboniferous

type.

Glossites amygdalinus and Sphenotus colus are also distinctly Carbon-

iferous species, but in their case the identification is admittedly doubtful.

I have not consciously made little of the evidence presented by Mr.

Butts, yet two of the species in his list are obviously identified with doubt

and it seems to me that of the remaining five the three invertebrates and

possibly also the two fishes belong to types in which the discrimination

of species is difficult and unsatisfactory even with very good material.

Furthermore, in interpreting the evidence I would think it wiser to ex-

tend the range downward of seven species rather than extend the range

upward of 52.

In seeking to determine whether the Bedford shale should be classed as

Devonian or Carboniferous, the problem is not perhaps whether it carries

a Chemung or a Kinderhook fauna; it is not a mere matter of correla-

tion, though correlation is involved. The fact seems to be that the Bed-

ford shale represents part of an interval between the base of the Missis-—

sippian and the top of the Devonian as those systems have usually been

GIRTY, GHOLOGIO AGH OF THE BEDFORD SHALE 309

defined and the real question is whether as such, from all the evidence

at hand, it belongs more properly with the one system or with the other.

The base of the Carboniferous system in this country, as usually recog-

nized, is the Kinderhook group of the Mississippi Valley; similarly the

top of the Devonian system is the Chemung group of New York. Now,

there is substantial evidence for believing that the Berea sandstone repre-

sents about the horizon of the Kinderhook and that it occurs several

hundred feet above the top of the true Chemung. The evidence for this

may be briefly summarized as follows: The representative of the Berea

in Crawford and Erie counties in northwestern Pennsylvania appears to

be the “Corry” sandstone. The “Corry” is more fossiliferous than the

Berea and contains a varied and characteristic fauna. The “Corry”

horizon carrying this fauna can be traced eastward to Cobhams Hill just

east of Warren, where it comes in immediately above what has been called

the “sub-Olean conglomerate” (Knapp formation), in the short interval

which separates that formation and the Olean conglomerate. Beyond

this the “Corry” horizon cannot be recognized, but it seems to be a mat-

ter of common agreement™ that the “sub-Olean conglomerate” at Warren

and the beds beneath represent the formations which in the Olean quad-

rangle come in below the Olean conglomerate, where an interval of about

500 feet, comprising the Oswayo and Cattaraugus formations, occurs

above the typical Chemung. Similar facts are indicated by I. C. White’s

work in Crawford and Erie counties,’* since he recognizes the Venango oil

sand group (which he calls Upper Chemung), with a thickness of 310

feet, and the Riceville shale with a thickness of 80 feet as intervening

between the Chemung proper and the formations for which he used

the names Corry and Cussewago. However many errors in detail there

may be in these tracings and correlations, it seems safe to conclude that.

an interval of 400 or 500 feet does intervene between the top of the true

Chemung and the “Corry” (Berea) sandstone in this area, which is prob-

ably represented in Ohio by the Bedford, Cleveland and Chagrin forma-

tions.

The first point to be considered in the paleontologic aspect of the

problem is the affinity of the Bedford fauna, its predominant Devonian

or Carboniferous facies interpreted on the facts of the general region in

which the Bedford shale and the Bedford fauna were developed.

Many genera and a few species, after a greater or less development in

the Devonian, pass upward into the Carboniferous, ranging to various

horizons in the Mississippian or even above. In most cases, there is no

-14 See the report by Glenn, Butts and Clarke already cited.

158 Sec. Geol. Sury. Pennsylvania, Rept. Q. 4, 1881.

310 ANNALS NEW YORK ACADEMY OF SCIENCES

general character or characters by which the Carboniferous species as a

whole differ from the Devonian species. In some instances, however,

certain general types within a genus appear to be restricted to one sys-

tem of rocks or the other. Thus, the punctatus group of Producti is a

distinctly Carboniferous development of the genus so far as known.

Now, the great bulk of the Bedford fauna, as will be seen from an ex-

amination of the table given on another page, belongs to types not char-

acteristic of either system. Most of them would not appear out of place

in either a distinctly Devonian or a distinctly Carboniferous fauna. In

such an association, one might say “This is a new species in this fauna”

but not “This is a Devonian species” or “a Carboniferous species,” as the

case might be. Thus most of the Bedford species, considered in their

broader relations, are ambiguous in deciding the Devonian or Carbon-

iferous affinities of an intermediate fauna. One might indeed take up

the Bedford fauna species by species and draw an inference from the

number of Devonian, of Carboniferous and of new species as to whether

the fauna should be grouped with the Devonian below or with the Car-

boniferous above. Such a careful canvass of the relationship of the dif-

ferent Bedford species would require more time than it has been possible

for me to give and would almost need be accompanied by a discussion of

each species, such as would be out of place in a paper of the present

scope. Besides this, as between closely related species in the Devonian

and the Carboniferous the conclusion reached in the identification would

many times be a matter of personal opinion. Comparisons sufficiently

ample have been made, however, to show that many of the Bedford spe-

cies are new and that the Carboniferous alliances are at least not more

numerous than the Devonian. J propose, on the other hand, to point

out a few instances of larger groups than species, about the identifica—

tion of which there can be less room for personal differences of opinion

and which, because they do represent larger groups, carry more weight

than species themselves, for I take it that the horizon which marked the

extinction of the genus Spirifer would be more noteworthy than that

which marked the extinction of some one species of Spirifer, such as

S. keokuk. Of such peculiarly pre-Carboniferous types, the first in my

list is the genus Pholidops, which has never the world over I believe been

found at horizons recognized as Carboniferous. The next on the list is

Delthyris, which has usually been identified in the Bedford fauna as

Spiriferina and which I have already discussed at some length. This is

a distinctly Devonian type of Spirifer and with one or two exceptions, to

which reference will be made later, has never been cited from Carbon-

iferous rocks. Next come the types which I have called Nucleospira ?

sp. and Camarospira ? sp.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE ant

These two forms present serious difficulties of exact identification,

being complicated with each other and with the two terebratuloids which

I have called Cranena aff. subelliptica and Cryptonella ?sp. These are

among the rarer forms of the Bedford fauna. When preserved in the

shale they are apt to be badly crushed, but they often retain the shell, so

that its structure can be determined. When preserved in the calcareous

nodules, the shell is not retained (or its structure is obscured), but the

proportions are not seriously altered. Thus, among these poorly charac-

terized, generally ovate forms there are clearly two types, one with a

punctate and one with a fibrous shell and of each type there appear to be

two species, distinguished more or less strongly by size and configuration.

Where the specimens have their real characters obscured by crushing or

in other ways (and this is true of many of them) they cannot be satis-

factorily placed in this scheme. The shells with fibrous structure have

the general appearance of Athyroids, and for such they might casually

be mistaken, but the ventral valve (and in one type both valves) is fur-

nished with a well-developed median septum. ‘This character is not only

alien to the Athyroids, such as Composita which the configuration sug-

gests, but I do not know of any Carboniferous genus which has at once

this shape and this structure. The larger of the two species suggests

Camarospira more than any other genus with which I am acquainted,

and the smaller more transverse one, which has a dorsal as well as a ven-

tral septum, is certainly very suggestive of Nucleosmra. I have even

observed what appear to be traces of fine sete on external molds.

It cannot be positively asserted that these forms belong to the genera

named, but it is true so far as I am aware that no genera having the

character of these Bedford shells are known in any Carboniferous rocks

of the Appalachian region. A few occurrences of Delthyris and Nucleo-

spira have been noted in the Kinderhook group of the Mississippi Valley,

but aside from this the Pholidops, the Delthyris, the Nucleospira ? and

the Camarospira ? are peculiarly Devonian types and are not found in

the Carboniferous.

On the other or Carboniferous side must be mentioned the Syringo-

thyris, which can, however, no longer be regarded as distinctly Carbon-

iferous in its generic range. The Bedford form is, however, identified

with a Carboniferous species. Again, I have a single very poor Spirifer

which seems to belong to the marionensis group (a Carboniferous type),

but which may be a somewhat abnormal S. disjunctus (a Devonian type).

Lastly there is a species of Pholadella which is more nearly allied to the

Carboniferous P. newberryi than to the Devonian P. radiata. These

Carboniferous affinities, it will be noted, are specific, while the Devonian

- ones are generic.

3479) ANNALS NEW YORK ACADEMY OF SCIENCES

To summarize the matter so far as considered, the Bedford fauna is

in many respects unique. It can be traced southward into Kentucky, but

it cannot be traced eastward into Pennsylvania. Its place in the “Brad-

fordian” of Pennsylvania has not been determined. It is distinct from

the “Bradfordian” fauna. It is distinct from the Chemung fauna. It

is quite distinct from the Chagrin fauna, which underlies it in the same

section, and which, while differing in important particulars from the

typical Chemung fauna, has nevertheless more of a Chemung aspect. It

is equally distinct from the overlying Berea (“Corry”) fauna, which has

more of a Mississippian aspect. It has a Devonian, or, as has sometimes

been said, a Hamilton facies, because, while it consists mostly of genera

which range into the Carboniferous and of species many of which have

Carboniferous affinities, it is nearly lacking in the strictly Carboniferous

types which abundantly accompany the latter at higher horizons and

proclaim the geologic age, and because it contains a few Devonian types

which very rarely and in the region under consideration never, so far as.

known, range up into the Carboniferous.

The Bedford and Cleveland formations may be lacking in northwestern

Pennsylvania owing to pre-Berea erosion, or to some other cause, but I

hardly believe this to be the case. If the Bedford does represent part of

the typical “Bradfordian” section, and if its fauna is a peculiar and local

development of the “Bradfordian fauna,” then we must enlarge the dis-

cussion to include the “Bradfordian” (“Upper Chemung’) faunas,

whose Devonian facies is conspicuous.*® In the one case (if properly

lying above the “Bradfordian”’ but removed by erosion in the typical! sec-

tion) the stratigraphic evidence, and in the other the paleontologic evi-

dence, is stronger for classifying the Bedford shale as Devonian. To

this must also be added the fact of a conspicuous faunal break between

the Bedford and the Berea, and the fact already noted that in the Berea

(“Corry”) we have, for the first time in this region, a fauna with a pre-

dominating Carboniferous aspect, one which shows many new features

when compared with the typical Mississippian, but which is distinguished

by the absence of most of the Devonian types of lower horizons and the

presence of many characteristic Carboniferous ones.

' Thus far it seems that the evidence has been strongly favorable to

classifying the Bedford with the Devonian. If we broaden the discus-

sion so as to include a larger field, that of the typical Mississippian area,

which apparently represents a conspicuously different province, or at

16 See the “Upper Chemung” species of Hall’s New York reports and Butts’s lists of

the faunas of the Knapp and Oswayo formations. These show a fauna with well

marked differences from the typical Chemung, yet with, in my opinion, a distinctly De-

vonian aspect.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 313

least represents a conspicuously different faunal development, the ques-

tion becomes more confusing and the conclusion somewhat less satis-

factory.

A search among the American faunas now known for one which is

comparable with that of the Bedford shale probably reveals none so simi-

lar in a general way as a certain phase of the Kinderhook developed at

Hamburg, Illinois, and the closely related one developed in the Glen

Park limestone member of the Kinderhook of Missouri. The constitu-

tion of these faunas, generically considered, is surprisingly close and

many of the species seem to be related. These may be arranged in

parallel columns, as follows:

Limestones of the Kinderhook at Glen

Sealine, Salle Park, Louisiana or Hamburg

Schuchertella herricki Schuchertella chemungensis

Productella pyxidata var. bedfordensis Producitella pyxidata

Rhipidomella n. sp. Rhipidomella missouriensis

Cryptonella ? sp. Cryptonella sp.

Delthyris n. sp. Delthyris missouriensis

Syringothyris carteri Syringothyris carteri

Spirifer aff. marionensis ? Spirifer marionensis

Athyris aff. hannibalensis Athyris hannibalensis

Macrodon hamiltonie Macrodon sulcatus

Leda diversa Leda diversoides

Bellerophon aft. jeffersonensis Bellerophon jeffersonensis

Platyceras sp. Platyceras erectoides

Tropidodiscus cyrtolites ? Tropidodiscus cyrtolites

There is also a Chonetes in both faunas, though of not very close rela-

tionship; a Camarotechia, though the Bedford form is large and the

Hamburg form small; and a Nucleospira, though also specifically dis-

tinct. In fact, while many of the same genera are present in both areas,

there are very few species which are really identical, and those for the

most part belong to genera in which the identification of species is diffi-

cult and a satisfactory identification impossible. Syringothyris carteri

is an example.

Professor Weller’” has pointed out the close relationship which exists

between the Kinderhook fauna at Glen Park, Missouri, and that from

the odlitic beds at Hamburg, Illinois, and he has also called attention to

the conspicuous Devonian facies which these faunas present (p. 463), a

resemblance which (like that of the Bedford) would seem to ally them

with the Hamilton rather than with the later Devonian faunas. Pro-

fessor Weller finds that 12 out of the 31 species at present known from

17 Acad. Sci. St. Louis, Trans., vol. 16, p. 462 et seq. 1906.

314 ANNALS NEW YORK ACADEMY OF, SCIENCES

Glen Park can be paralleled in the Hamilton formation of New York,

while only six species have parallel forms in the Chouteau limestone of

Missouri. Although the Chouteau fauna is so near at hand, he is able

to find specific identity in only two species and those are Schuchertella

chemungensis and Tropidodiscus cyrtolites. When we consider this fact

and that the fauna comprises such Devonian genera as Hunella, Atrypa,

Nucleospira and Delthyris, not to mention the fish Ptyctodus eastmani,

it would seem that the Devonian proclivities of the fauna far outweigh

the Carboniferous ones, even with due consideration for the two doubt-

- fully identified crinoid genera. This evidence is largely neutralized,

however, when other factors are taken into consideration.

The Kinderhook faunas of the upper Mississippi Valley show local

' facies to an almost unprecedented degree. To some extent, this differ-

entiation may have a zonal explanation, but it is also probably local and

environmental, since the lithologic character of the beds is also extremely

variable. Professor Weller recognizes a northern and southern type of

Kinderhook fauna which were contemporaneous, but almost entirely dif-

ferent. The Chouteau limestone exemplifies the southern fauna and

with this, as just noted, the Glen Park fauna has only two species in

common, although Professor Weller apparently regards them as occupy-

ing the same horizon. Both the northern and the southern faunas are

also highly diversified.

Beneath the fauna of the odlite at ghasiues, referred to above, which

so closely resembles that of the Glen Park limestone, there is another

having a considerably different facies. The latter Professor Weller cor-

relates with the well-known fauna of the Louisiana limestone and this

in turn with the typical Kinderhook of that ilk, which corresponds to

the lower and larger portion of the Kinderhook section at Burlington,

On the other hand, he correlates the fauna of the odlite at Hamburg

with the Glen Park fauna and the Glen Park fauna with the Chouteau

fauna, and with the upper part of the Kinderhook section at Burlington,

if I understand him aright.

Several very different facies are presented by these faunas. That of

the Louisiana limestone (at Louisiana, Missouri, and Hamburg, Illinois)

is distinctly more Carboniferous than that of the odlitic limestone at

Hamburg and Glen Park, which, as already noted, are rather conspic-

uously Devonian, though they occur above the other in stratigraphic

position. The faunas of the Chouteau limestone and the topmost Kin-

derhook at Burlington, with which the faunas of the odlitic limestone at

Glen Park and Hamburg appear to correlate, are still more conspicuously

Carboniferous, and they have so been recognized for a long time.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 315

As I have just shown, the fauna of the Bedford shale and the fauna of

the odlitic limestone at Glen Park and at Hamburg are in some respects

strikingly alike, but, though the resemblances are undoubted, there are

also numerous and important differences. The resemblances consist of

the presence in both faunas of identical genera and of related species.

Identical species, however, are few and not of the first importance. The

Delthyris, the Nucleospira, the Macrodon, etc., of the Bedford shale are

not the same species as the Delthyris, the Nucleospira, the Macrodon,

etc., of the odlite at Glen Park and at Hamburg. Correlation by similar

species is certainly much more hazardous and less satisfactory than corre-

lation by identical species. Indeed, although we of course know that all

these faunas are more recent than the Hamilton, the table compiled by

Professor Weller would indicate that the Glen Park fauna is almost as

closely related to the Hamilton faunas as it is to that of the Bedford

shale and much more closely related to the Hamilton than to the contem-

poraneous Chouteau fauna. Restricted to their own showing, therefore,

I believe that a correlation of the Bedford and Glen Park faunas would

not be justified, except in a very provisional and tentative manner.

However that may be, if, instead of considering the two faunas as iso-

lated occurrences, we include, as we are forced to do, the faunas asso-

ciated or correlated with them—the typical Kinderhook, the Chouteau

and the Louisiana faunas of the Mississippi Valley in the one case, and

the “Bradfordian” faunas of Pennsylvania in the other—it seems clear

that we have two entirely distinct faunas, the one showing a strongly

Carboniferous and the other a strongly Devonian facies, and we cannot

conclude that they are contemporaneous sees of the same faunal

zone on any evidence now known.

The Bedford and Cleveland shales cannot be definitely identified in

the “Bradfordian” rocks of northwestern Pennsylvania, either litho-

logically or paleontologically, but there is an interval between the Berea

(“Corry”) sandstone and the Venango oil sand group which seems to

correspond in a general way to that represented in Ohio by these forma-

tions, and I personally but little doubt that Bedford and Cleveland do

correspond to strata in the “Bradfordian.” Even, however, if they do

not, and the “Bradfordian” with its strongly Devonian fauna does en-

tirely underlie the Bedford, I believe that the correlation of the Bedford

shale with the odlites at Hamburg and Glen Park would not be justified

at present.

The small number of identical species and the almost complete ab-

sence of all those characteristic Carboniferous types which by Professor

Weller’s correlations occur at the same horizon as the odlitic limestones

316 ANNALS NEW YORK ACADEMY OF SCIENCES

at Glen Park and Hamburg or below, though not yet found associated

with them, would, in the absence of substantial evidence to the contrary,

indicate that the Bedford shale was not really a contemporaneous forma-

tion.

The correlation of other horizons in the same sections is intimately

connected with that of the Bedford shale. Professor Weller?® recognizes

in the famous goniatite-bearing limestone at Rockford, Ind., a repre-

sentative of the Chouteau limestone or southern Kinderhook. It is

perhaps impossible to tell in fact, as it is certainly impossible to tell from

his discussion, whether the limestone at Rockford corresponds to that

part of the Chouteau which correlates with the northern Kinderhook or

to that part which he recognizes in the Burlington section as occurring

above the northern Kinderhook. If the Bedford represents the Glen Park

horizon, it would apparently on the one hypothesis correlate with the

goniatite-bearing bed at Rockford (and it does contain some goniatites

of scarcely determinable genera), while on the other hypothesis it would

come in above it. The position of the Bedford shale above the black

Cleveland shale is at first suggestive of the position of the goniatite-

bearing bed at Rockford above the Devonian black shale of Indiana, but

there is no assurance whatsoever that the two black shales represent the

same horizon and, even if such were shown to be true, it would not neces-

sarily follow that the succeeding formation in the one case corresponded

to the succeeding formation in the other. Indeed, until quite recently

it has been the general consensus of opinion that the goniatite-bearing

bed and the black shale beneath were quite separate and distinct forma-

tions divided by a long-time interval, the one of Carbonifrous, the other

of Devonian age, and no satisfactory evidence has yet been produced for

believing otherwise. On the other hand, such facts as I am acquainted

with both of stratigraphy and paleontology go to show that the Bedford

and Cleveland shales are related in the closest manner and must be

classed together wherever they are classed.

The early Mississippian sections of Ohio and of the Mississippi Valley

show great differences of development, both in the sediments which

accumulated there and in the animal life which those sediments helped

to condition. They probably constitute distinct provinces. There are

no faunas in Ohio closely allied to the typical Burlington and Keokuk

faunas,—nothing to correspond to the rich development of crinoid life

which is found in those faunas and which doubtless did much to deter-

mine the character of the associated life, unless still different influences

determined both. The 18 species of crinoids known from the Cuyahoga

18 Loc. cit., p. 469.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 317

shale, though belonging to genera well represented in the early Missis-

sippian of the Mississippi Valley, do not occur outside the State. Any-

one, however, who will compare the fauna of the Chouteau limestone

with that of the Cuyahoga shale, as found at such points as Medina,

Richfield, Lodi and Royalton, cannot fail to find great similarity and not

a few identical species. I am not prepared to state the exact extent of

this resemblance, but my studies would indicate strongly that, if the

Cuyahoga fauna is to be found anywhere in the Mississippi Valley, it is

to be found in the Chouteau hmestone. The Waverly localities which I

have mentioned are all, I believe, in the upper Cuyahoga. By definition,

the Chouteau limestone is part of the Kinderhook group and therefore

in stratigraphic position inferior to the Burlington limestone, but I am

much disposed to think that the Chouteau limestone really correlates

with the lower Burlington, the fauna which we know as the Burlington

fauna being developed in and largely confined to the upper Burlington.

I have already given a list of the fossils found in the Berea (“Corry”)

sandstone which underlies the Cuyahoga shale. This fauna contrasts

_ strongly with both the Bedford fauna below and the Cuyahoga fauna

above. It has a much more marked Carboniferous aspect than the Bed-

ford fauna, even if we exclude the faunas apparently contemporaneous

with the Bedford having a more distinctly Devonian facies. Though,

of course, showing great individuality, the “Corry” fauna is not only

distinctly Carboniferous, but in some of its elements it is distinctly

Kinderhook, as for instance in the genus Paraphorhynchus, a type which

Professor Weller regards of special importance and which is said to be

characteristic of the northern Kinderhook.

_ It is interesting to find three faunas in the Ohio section showing re-

semblances, more or less illusory perhaps, to these three aspects of the

Kinderhook faunas of the Mississippi Valley, and it is also interesting to

compare the stratigraphic relations of these faunas in the two areas on

the assumption that the Cuyahoga shale correlates with the Chouteau

limestone, the Berea sandstone with the northern Kinderhook (the

Chonopectus fauna of the Kinderhook sections at Burlington), and

the Bedford shale with the odlite at Glen Park or Hamburg, as is to

some extent suggested by faunal similarities. According to their strati-

graphic relationship in typical sections in the Mississippi Valley, the

Bedford shale should not lie below the Berea sandstone, but above it. It

should in fact even be contemporaneous with part, if not with all of the

Cuyahoga shale. Tf, however, the Kinderhook relationship of the Bed-

ford be eliminated, as I believe it can be eliminated owing to its probable

relationship to other “Bradfordian” faunas, this contradiction largely

318 ANNALS NEW YORK ACADEMY OF SCIENCES

disappears. The Cuyahoga and Berea together represent the Kinder-

hook of the Burlington section (in which, if I understand him aright,

Professor Weller thinks that the upper 15 feet corresponds to the upper

part of the Chouteau, both stratigraphially and faunally, while the lower

part of the two sections corresponds stratigraphically but not faunally),

and they have the same relative position in both sections. If my hypoth-

esis of the equivalence of the Chouteau with the lower Burlington is

correct, then of course the Berea alone represents the entire Kinderhook

section at Burlington and presumably its correlates at Louisiana and at

Hamburg with their varying faunas.

Professor Weller, as already described, recognizes two types of Kin-

derhook faunas, the northern one, the typical Kinderhook, not being

found at all southward in southern Missouri and Arkansas; the other,

the Chouteau, occurring in Arkansas and Missouri and represented by

a few feet of rocks above the northern Kinderhook in the section at Bur-

lington. Professor Weller’s interpretation of these facts is that the two

faunas were developed contemporaneously in disconnected basins, to the

more northern of which the southern fauna gained access near the close

of Kinderhook time. Tentatively, I would prefer to explain these rela-

tions by supposing that the southern Kinderhook was entirely later than

the northern and was represented in the Burlington section not by the

topmost Kinderhook alone, but by the lower Burlington also. However

that may be, the disappearance southward of the northern Kinderhook

fauna is somewhat suggestive of the southward thinning of the Bedford

and the Berea formations, as recently described by W. C. Morse and

A. F. Foerste.*®

The careful stratigraphic work of these writers, combined with that of

Professor Prosser, indicates that the Bedford and Berea gradually thin

to a feather edge and presumably disappear as recognizable formations

in east-central Kentucky. To some extent, they also lose their distinct-

ive lithologic characters, so that Morse denominates as “Bedford-Berea”

the interval of shale and sandstone which they fill between the black

Ohio and Sunbury shales. The tracing by stratigraphy is corroborated

by the occurrence of more or less characteristic Bedford fossils at the base

of this interval in Kentucky, and Dr. Foerste himself very justly raises

the question whether the final appearance of these sediments, which con-

sist of shale alone, should not be referred solely to the Bedford forma-

tion, and the preceding occurrences in which the shales predominate

below and the sandstone above should not be divided into Bedford and

79 Jour. Geol., vol. 17, p. 164 ét seq. 1909.

GIRTY, GEOLOGIC AGE OF THE BEDFORD SHALE 319

Berea, respectively. In view of the fact that the Berea possesses a char-

acteristic fauna which has not been found in the sections under consid-

_ eration, whereas the Bedford fauna has been found there, and in a shale,

and at the base of the interval, it seems to me that that portion of the

interval would better be identified as Bedford alone, whatever is done

with the upper part, though the evidence would suggest the advisability

of calling the upper sandy beds Berea where they are present, and the

whole interval Bedford where they are not.

The bearing which this aspect of the Bedford-Berea stratigraphy has

on the question of the geologic age of the Bedford shale is not entirely

clear. From one point of view, one might say that it did not affect the

classification of the beds at all, except insofar as it made them difficult

to distinguish in the field and to delineate on a map. On the other

hand, it might be urged with some force that since by the expansion and

differentiation of the Chattanooga shale in a northward direction, that

formation seems to cover an interval including the lower Cuyahoga. the

Berea, the Bedford, the Chagrin and probably the Huron formations,

and in a manner to bind them together into one group of sediments,

they ought all to be classed as Devonian or all as Carboniferous. This,

however, does not at all agree with the facts, where these formations are

differentiated and developed in an unequivocal manner, and I believe

that it should not prejudice such a classification of the rocks as is indi-

cated by the facts ascertainable under those conditions.

Therefore, while the weight of the evidence is not entirely cast on

one side of the question, I believe that so far as the facts are known

they indicate the line at the base of the Berea sandstone as the proper

position of the Devonian-Carboniferous boundary in northern Ohio.

This is because that boundary is marked by an unconformity, by the

presence of a basal sandstone and by a pronounced faunal change, such

that while the fauna of the Berea (“Corry”) sandstone has a distinctly

Carboniferous facies and is probably to be correlated with the Kinder-

hook group of the Mississippi Valley, that of the Bedford shale, though

its stratigraphic position is above the typical Chemung, has, in connec-

tion with the other “Bradfordian” faunas, a distinctly Devonian facies.

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CHANGES IN THE BEHAVIOR OF THE EEL

DURING TRANSFORMATION

BasHFORD DEAN :

NEW YORK :

PUBLISHED BY THE ACADEMY

5 DrcemBer, 1912

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum oF Narurau History, 1817-1876) . —

OFFICERS, 1912

President—HMrrson McMIttin, 40 Wall Street

Vice-Presidents—J. E>pMUND WoopMAN, FREDERIC A. Lucas

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[Annats N. Y. Acap. Scr, Vol XXII, pp. 321-326, 5 December, 1912]

CHANGES IN THE BEHAVIOR OF THE EEL DURING

TRANSFORMATION

By BasHrorp DEAN

(Read in abstract before the Academy 13 May, 1912)

The literature of animal behavior gives as yet little attention to the

changes which occur in animals during the period of metamorphosis.

This is a gap in our knowledge not remarkable perhaps when we con-

sider how little is yet recorded of the behavior of many types of adult

animals, even of common forms. None the less, it is precisely during

the period of transformation that one may expect to find clues as to

interesting conditions in mind-mechanism, for during this short period

adjustments are completed which change, as it were, one functional

“species” into another; for an animal may remain for years in its larval

form almost unaltered, and it may subside again into a changeless form

after a kaleidoscopic transformation. In fact, the more sudden the

change in transformation, the more interesting it should be from the

point of view of connecting habits with structures, for it would here

bring into sharpest relief morphological changes and make them the

more easily linked with changes in behavior.

In the larval history of fishes, observations in this field have rarely

been recorded. The teleosts, where conspicuous larval stages occur, are

little studied, even in the case of those members of the group which have

the most complete metamorphosis.

The form-changes of eels have been described by a number of authors

(Grassi, Calandruccio, Cunningham, Eigenmann and others)., and the

changes are so marked that we can readily predict from them striking

changes in behavior. That the latter actually occur, and in marked

degree, was clearly brought home to the writer when an opportunity

came to him in Japan (Misaki) to observe the transformation of a

Leptocephalus into a Conger,—possibly Conger (Leptocephalus) mala-

baricus (Day).* His notes, especially upon its behavior, are perhaps

worthy to be recorded on account of the interesting nature of the “larva”

and from the fact that this form is not apt to be observed. In point of

fact Leptocephalus seems rarely to have been kept living in an aquarium

tWRANCIS DAy: “The Fishes of Malabar.” Pl. xix. 1865,

(321)

322 ANNALS NEW YORK ACADEMY OF SCIENCES

more than a few hours. The specimen in question, I may mention, was

in perfect condition when taken. It was noticed in the bag of a seine,—

by accident rather than by design,—for had the fish not been actively

moving at a particular moment, it would have escaped unnoticed on

account of its glassy transparency. It proved to be hardy and lived in

an aquarium for over three weeks, during this time undereot its

metamorphosis.

September 13. Larva (Fig. 1) almost colorless, even in light of dif-

ferent intensities; it is rarely at rest; it is apt to swim rapidly and with

Fic. 1.—#el larva. September 13. About natural size

a kind of lurching movement. When advancing slowly, its height, which

insures contact with a large surface of water, allows it to move with

precision,—in the sense that a pencil held vertically in the first in-bent

curve of the fish’s body will not be touched by the fish as it advances,—

in other words, that the fish does not show lost or slipping movements.

When resting, the young fish arranges itself

in irregular vertical coils, thus probably

keeping its balance. When disturbed

(snout touched with a pencil point), the

fish retreats tail foremost, the head remain-

ing passive: if disturbed again, the head

will be quietly drawn back, the motion

starting as before with a withdrawal of the

tail and hinder trunk. If disturbed repeat-

Fic. 2.-—Hel larva. Retracted edly, however, the fish will either swim

meee about actively or draw itself into a close

coil (Fig. 2). This position, however, it will sometimes assume without

artificial stimulus, e. g., after it has become “tired” swimming around

the wall of a circular jar. Leptocephalus from time to time secretes con-

siderable mucous: this remains attached but is finally “brushed off’ (in

a mass) at the tail end of the body. Such a bit of slime will occasionally

be touched by the young eel when swimming about; it is evidently dis-

tasteful, for the young fish speedily frees itself, shaking its head in a

curiously energetic way.

DEAN, BEHAVIOR OF THE EEL DURING TRANSFORMATION 3923

September 15. The fish is now more easily seen. There is a slight

clouding of its transparent sides, especially near the lateral line (Fig. 3).

Its eyes are conspicuous and show numerous movements. It is more

active than in the earlier stage, sometimes swimming with broad undu-

lations (Fig. 4) different in type from earlier movements. A patch of

ys WH ddddiee

j DN \\ ° \\

Fic. 3.—£el larva. September 15

color, brownish orange, appears on the ventral body wall, just behind the

gill opening (jugular villi).

. September 18. The larval length and breadth are rapidly becoming

reduced (Fig. 5). The caudal fin and dorsal ridge appear. The colored

jugular patch has now de-

veloped into a velvety mass bd

of lighter color, and possibly

serves as a larval adhesive

organ, which hangs freely in

the water. The intestine

can be outlined. Pigmenta-

tion is noted, especially

along the lateral line and on

the head-roof, and the en-

tire fish has a faint purplish , \

tone. It remains more often

at the surface than before, \\y eee

here occasionally floating

and swimming on its side,

now and then thrusting its

head out of water. It remains longer in one position than heretofore.

If disturbed (head touched), it will wriggle its head backward,—and

does not initiate the backward movement from the tail as in the earliest

stage.

September 19. Changes progress rapidly (Fig. 6). The coloration

is distinctly purple, with whitish spots near the tail, and pigment patches

on the ventral wall of the head.and within the neural axis. Vertebre

lie. 4.—Hel larva. Position in swimming

394 ANNALS NEW YORK ACADEMY OF SCIENCES

Fie. 5.—Eel larva. September 18

Fic. 6.—LHel larva. September 19

Fic. 7.—Hel larva. September 22

Fic. 8.—#eil larva. September 28

Fic. 9.—Eel larva. October 2

Figures about natural size

DEAN, BEHAVIOR OF THE EEL DURING TRANSFORMATION 325

are now seen. The patch of jugular villi is reduced in size. The be-

havior of the young fish is eel-like. It remains motionless for longer

periods, occasionally lifting and turning its head, and there are pro-

nounced movements of its opercula.

September 22. Larval coloration is apparent in the white spots above

and below the tail. In general the advances are clearly in the direction

of the mature eel. The vertebre are conspicuous; the visceral wall be-

comes opaque; the jugular larval organ is represented by a clump of

scattered filaments; the gill region is more conspicuous (Fig. 7); the

gill arches show the red lamelle, expand broadly and contract; the

mouth opens wide; there is no movement in the neck region; the pec-

toral fins function; and the swimming is snake-like, with more effort

than propulsion, 1. ¢., slipping, unlike the precise movements of earlier

stages.

September 26. From now onward, the changes are less noticeable.

Larval coloration is retained, e. g., in the light colored spots. The trunk

is opaque, even in the gill region.

September 28. At this stage, the last trace of the larval jugular organ

was noticed. The body is thickened; the white spots have disappeared

(Fig. 8).

October 2. The last stage recorded (Fig. 9). Transformation is

practically complete. Measurements of this contrasted with the earliest

stage show a surprising shrinkage in the length and height of the young

eel,—more exaggerated even, than in the cases described by Grassi. In

the present instance, the young fish is about one-half the length of the

earliest stage, and one-third of its height, after a growth period of about

three weeks. It is another example of the paradox that development

may be accompanied by considerable diminution in size.

Especially interesting in the foregoing transformation is the rapidity

with which the behavior of the young eel changes. This is not brought

out in adequate detail in the present note, but it may be said that the

observer could not but feel that the larva behaved like an animal suite

of a different species from the one of the days before, or of the days

following. This state of affairs predicates, obviously, kaleidoscopic

changes in elements of the central nervous system, and astoundingly

delicate and rapid adjustments; but whether these can be actually de-

termined, 7. e., in the physical characters of the cells of brain and cord,

must yet remain an open question. It can be solved only when an

abundant material of Leptocephalus falls into the hands of a specialist

who can bring to his aid the latest neurological technique.

326 ANNALS NEW YORK ACADEMY OF SCIENCES

From a phyletic point of view, on the other hand, the origin of the

rapidly changing behavior correlated with morphogenetic changes is less

difficult to understand. We have, first of all, numerous grounds for

concluding that the larval stages of teleosts are secondary, and that

eccentric forms,—i. e, those with extraordinary fins, colors, outlines,—

are derived from “larve” in which such extreme structures did not exist.

‘In the case of the eels, therefore, we can reasonably picture a progressive

form of development, such, for example, as occurs in many of the older

groups of teleosts. We next suggest that within this progressive series

of closely similar stages, one stage should become especially important

as adapting the young to a particular environment. The young eel then

would tend to remain longer unchanged in the special environment

favorable to its feeding, movements, lack of pigment, temperature-re-

quirements, etc., and this phase in its life-history would come to sup-

plant the adjacent steps in the progressive series. In other words, if we

erant that the development of a young eel (the montée) might be accom-

plished in the space of fifty weeks, and that at the end of this period it

completed the fiftieth of its intergrading stages, we could also admit

that with the same total period of larval growth certain of the stages

might have expanded while others contracted. Thus, to take an ex-

ample, stage twenty, which earlier may have been passed through in a

week, might become successively protracted to two weeks, three weeks,

or months. And the stages of the montéc intervening between twenty

and fifty would be correspondingly reduced. The interval was at first

thirty weeks, and included thirty stages; it was next, say, fifteen.weeks

in which to represent thirty stages, and finally in the case of our Lepto-

cephalus, it was reduced to the astonishingly short term of three weeks,

in which to represent the many stages. In such an instance there can

be little question that the marked changes in behavior are correlated

with abbreviated phases of development.

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THE NEW YORK ACADEMY OF SCIENCES

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OFFICERS, 1912

President—BKMERSON McMILtin, 40 Wall Street

Vice-Presidents—J. EDMUND WoopMAN, FREDERIC A. Lucas

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[Annats N. Y. Acap. Sct., Vol. XXII, pp. 327-333. 20 December, 1912]

NOTES ON THE HABITS OF A CLIMBING CATFISH (ARGES

MARMORATUS) FROM THE REPUBLIC

OF COLOMBIA

By R. D. O. Jounson

(Read in abstract before the Academy 13 May, 1912)

Introductory remarks, offered by BASHFORD DEAN at the meeting—The

group of catfishes (Siluroids) holds a puzzling place among fishes. That it

represents one of the ancient groups of bony fishes, there can be no doubt, but

whether Siluroids are descended from some special line of ganoids or whether

they have been derived through a long series of specializations from some

ancestor essentially carp-like remains ever an open question. The trend of

later work, certainly, tends to ally them more closely with plectospondylous

forms, but many of their most important structural characters have never

been explained on such a basis.

Thus, the limb-girdles of some of the catfishes, with their accompanying

muscles, have appeared to be primitive, and there has, as far as I am aware,

been adduced no evidence to show that these structures were derived from

highly specialized conditions of such living plectospondyls, for example, as

characinids. The habits of Siluroids, which would help to explain the signifi-

cance of these abdominal structures, have not been known to be remarkable,

and there is no suggestion, therefore, that the characters in question might

but be interpreted as highly modified rather than primitive. Accordingly, the

present paper of Mr. R. D. O. Johnson merits, I believe, the attention of the

Academy, for he shows that under conditions of stress, the ventral structures

of the catfish Arges have an especial value to the fish in enabling it to creep

against the strongest currents and to climb with great rapidity and skill. The

conclusion, therefore, is evident that we may now reasonably interpret the

puzzling fin-structure of Siluroids as developed in relatively recent times, and

as having little significance in terms of more ancient groups.

Mr. Johnson, it may be mentioned, spent several years in the highlands of

the Republic of Colombia, and although the region he studied has been visited

by but few naturalists, it is nevertheless hardly to the credit of our “cloth”

that these observations on fishes should first be made by a mining engineer.

The creeks and rivers of the Andes Mountains in the Republic of

Colombia, South America, are torrential in character. The great major-

ity of them are but a succession of falls, cascades, pot-holes and short

“rifles.” The rainfall in the mountains is heavy and the rock under-

lying the stream beds is schistose in character and comparatively soft.

whe rate of erosion is exceedingly rapid, yet the grade lines of these

(327)

598 ANNALS NEW YORK ACADEMY OF SCIENCES

streams stand at high angles. This unstable condition seems to be due

to the elevation of the Andes during a late geological period. The heavy

rainfall, at times amounting to four or five inches within a few hours,

produces floods of immense volume. These go charging down the can-

yons with fearful fury, and at times it would appear that nothing could

withstand their sweeping energy. Yet these turbulent waters are the

habitat of fishes so wonderfully adapted to their surroundings that they

are able to grow and to multiply in great numbers.

In external appearance they resemble the catfish or horned pout of the

north. The skin is smooth and scaleless. The color is a dark mottled

gray shading into a slightly yellowish tint on the posterior parts. They

rarely attain a length greater than twelve inches. As an article of food

Fic. 1.—Arges marmoratus Regan ; side view

they are esteemed by the natives and are well known by the local name

“Capitan.” They have lately been described by C. Tate Regan as Arges

marmoratus.*

Under usual conditions they are clumsy and awkward swimmers, wrig-

gling through the water like tadpoles, but as creepers and climbers they

are without rival in the fish family. The mouth is small, but is sur-

rounded by a broad, soft, rubber-like flap, very thin and flexible at the

edges (Fig. 2). It is a sucker mouth and the entire mechanism is so

perfectly adapted to the needs of the fish that it finds no difficulty in

firmly attaching itself to any convenient object. It is this ability to

make a quick anchorage that enables the fish to stay at home when nature

seems bent upon sweeping the canyons and water-courses clear of every-

thing movable.

If, however, these fish were able only to keep themselves from being

washed out in flood times, they would be insufficiently equipped to main-

tain an existence in these mountain streams. If they depended upon

their imperfect swimming alone as a means of locomotion, whatever

migratory movement they attempted would inevitably have to be made

1 Trans. Zodlogical Society of London, XVII, p. 314. 1904.

JOHNSON, HABITS OF A CLIMBING CATFISH 399

m a down-stream direction. The final result would be the same as

though they were unprovided with a means of anchoring themselves at

will. But they are equipped with another and very efficient apparatus

for locomotion, The flat sucker mouth is half of the mechanism; the

other half is located on the belly. Under the skin of the ventral side,

just behind a line joining the pectoral fins, there is a triangular bony

plate to which are attached the ventral fins (Fig. 2). The main anterior

ITuscles

Fig. 2.—Arges marmoratus Regan; ventral view

ribs of these fins are broad and flattened, and the flat. surfaces are thickly

studded with small, sharp teeth pointing backwards. The triangular

plate and its attached fins are free to move in a longitudinal direction

through a distance equal to about one-sixth of the length of the fish.

This movement is accomplished by means of four muscles in two pairs

attached to the plate; the anterior pair extending from their attachments

on each side of the plate forward to the middle point on the bony arch

just below the gill openings; the posterior pair extending from an

330 ANNALS NEW YORK ACADEMY OF SCIENCES

attachment at the center of the posterior edge of the plate to the anal fin.

It is evident that the fish is able to create a suction pressure in the re-

gion of the plate, though how this is accomplished is not apparent from

the structure.

By means of the alternate action of the mouth and of this curious

apparatus, the fish is able to creep against a current that would baffle

its efforts entirely, if it relied alone upon its fins and tail. When it

Orifices for the

Inflow of water

to Gills

—--- rr

lig. 3.---Arges marmoratus Regan; dorsal view

is engaged in creeping or in sticking fast to some object, the sucker

mouth necessarily is closed. It is evident that the gills must be sup-

plied with the life-maintaining flow of water through some other avenue.

At the upper extremity of each gill slit there is an orifice provided with

a valve opening inward (Figs. 1 and 3). During the diastole of the gill

covers, the water flows inward through the orifices and is expelled

through the gill slits during the systole. |

JOHNSON, HABITS OF A CLIMBING CATFISH 331

On clear sunshiny days, these fish may be seen in the depths of the

clear water hitching themselves along over the surfaces of rocks, occa-

sionally swimming short distances in the more quiescent places, but

seeming to depend for locomotion primarily upon their creeping mechan-

cays --as

“i

= ENeN

SANS SS a

\ SS =o

SS =:

! ! i Al

iit

\ZAI e

ye — a

) : sy Wy

ay ; maT = =

| AWS es il

ia yj PAZANMNY, \“

MMC ZZ

LEE

Fic. 4.—Section of a pot-hole, twenty-two feet deep, in Santa Rita Creek, Colombia,

showing ‘‘capitanes’’ ascending its rocky walls

ism. They are to be found in all parts of these mountain streams, from

the most slender tributaries to the foot of the mountains. It is evident

from this fact that they are able to travel up stream. They are too

539 ANNALS NEW. YORK ACADEMY OF SCIENCES

‘sluggish in movement and are provided with a swimming apparatus

-altogether too inefficient to enable them to dash up the high and frothy

falls.

At one time, the writer had occasion to divert the water of a small

mountain stream so that access could be gained to a deep pot-hole from

which the water, rock and gravel were subsequently removed. This pot-

hole was twenty-two feet deep, nearly circular in horizontal cross-section

and it varied in diameter from six to ten feet. Generally, the sides ap-

proached the vertical and in some parts inclined inwards. When the

water had been lowered to within four feet of the bottom, the remaining

water was seen to contain a large number of “capitanes.” ‘They were

greatly excited and distressed and were swimming and creeping about

in all directions. A small stream of water in a thin film ran down one

side of the pot-hole from a leak in the dam above. Several fish, after

nosing around the edge of the water, discovered this small inflowing

stream and started to creep up in it, but becoming frightened by the

movements of the working men near, dropped back. When work was

stopped for the noon hour, four of the smaller fish started up, following

the thin stream of water. The water ran over their noses, down their

backs and trickled off their tails in small streams. They would hitch

themselves up rapidly for the distance of a foot or so and remain quiet

for a minute or two; then another foot and another rest. In half an

hour, the four had reached the water in the pool at the foot of the dam

above. In making the ascent, they were obliged to pass a part of the

wall, about two feet in length, that inclined inward at an angle of about

30° from the vertical. When they reached this overhanging part, in no

observable manner did they change their tactics, but they ascended it as

rapidly and safely, and apparently with no more effort than the other

portion of the wall. During the afternoon, several more of the fish

climbed out. A large number were in the water at the bottom of the

hole when work was suspended for the evening. In the morning not a

fish remained.

For the greater part, the path followed by the fish in making their

ascent lay over smooth, water-worn surfaces free from any coating of

vegetable matter. The upper part, however, was covered by a thin film

of an alga-like growth that may have served for the engagement of the

‘sharp-pointed teeth on the movable ventral fins. The total vertical dis-

tance through which the fish climbed measured eighteen feet. When

‘undisturbed, they covered the distance without a slip or fall. The water,

diverted around this pot-hole, flowed through a large pipe and fell from

the end upon the steeply inclined water-worn rock at the side of the

JOHNSON, HABITS OF A CLIMBING CATFISH 333

channel below. A day or two after the water had thus been diverted, a

dozen or more of these fish were observed to be clinging to the rock at

the foot of the fall at the end of the pipe. They were evidently on their

way up stream, but had encountered an artificial condition that inter-

rupted their further progress. They were nosing about in search of a

small stream or film of water sufficient to keep their gills wet and to lead

them to the main body of water above. As there was no such stream,

their further progress was prevented. ‘They made no observed attempt

to swim up the fall, but confined their efforts to making short excursions

up the rock above the water. Failing to find any leading stream, they

crept back.

They deposit their eggs in the deepest pot-holes and attach them indi-

vidually to the under sides of large rocks.

BLICATIONS

or THE

Ww YORK ACADEMY OF SCIENCES

(Lyceum OF N ATURAL HIsTory, 1817-1876)

are

ublications of the Academy consist of two series, viz.:

n tings and similar matter.

% ume of the Annals coincides in general with the calendar year

. old at the uniform price of three dollars per volume. The articles

“buted | in bundles on an average of three per year. The Bee of

rate articles depends upon their length and:the number of illus-

ns, and may be learned upon application to the Librarian of the

The author receives his separates as soon as his paper has

> i hed date of issue ae above the title of each pee

‘a are sent free to Fellows and Active Members. The

THs LIBRARIAN,

New York Academy of Sciences,

care of

American Museum of Natural History,

New York, N.Y.

THE NEW YORK ACADEMY OF SCIENCES |,

(Lycrum or NaturaL History, 1817-1876)

OFFICERS, 1912

President—EMeErson McMitui1n, 40 Wall Street y

Vice-Presidents—J. EDMUND WoopMAN, F'REDERIC A. Lucas

CHARLES LANE Poor, R. 8. WoopworrH

Corresponding Secretary—Hernry W. Crampton, American Museum

Recording Secretary—Epmunp Ot1s Hovey, American Museum”

Treasurer—HeEnry L. DoHERTY, 60 Wall Street |

Librarian—Ratreu W. Tower, American Museum

Editor—Epmunp Otis Hovey, American Museum ~

SECTION OF GEOLOGY AND MINERALOGY

Chairman—J . E. Woopman, N. Y. University

Secretary—Cuartes P. Berkey Columbia University

SECTION OF BIOLOGY

Chairman—FReEDERIC A. Lucas, American Museum

Secretary—Witu1am K. Gregory, American Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—Cuar.Es LANE Poor, Columbia University

Secretary—F. M. Peprrsen, College of the City of New York

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. S. WoopwortH, Columbia University

Secretary—FREDERIC LYMAN WELLS, Columbia University

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. Sct., Vol. XXII, pp. 335-337, Pll. XXXV-XXXIX.

27 December, 1912]

THE KINGSTON, N. M., SIDERITE!

By Epmunp Otis Hovey

(Read before the Academy 4 November, 1912)

In the year 1891, a prospector was ransacking the region along the

North Branch of Percha Creek, near the Solitary Mine, about four miles

north of Kingston, Sierra Co., New Mexico, in a search for horn silver

(cerargyrite), when he stumbled upon what he supposed to be a solid

mass of the ore for which he was hunting. ‘This was lying upon a ledge

of granite and, according to the account of the finder, had been brought

to view by erosion. The approximate position of the locality is latitude

32° 58’ North, longitude 107° 50’ West. A small fragment was broken

off and sent to an assayer, who informed the finder as to the real nature

of his specimen. Another small piece was sawed from the other end of

the mass. These two pieces would weigh together about 5 ozs. (142

gm.), while the mass as delivered to the Foote Mineral Co. weighed

28 lbs., 6 ozs. (12,870 gm.) ; hence the total original weight was about

28 lbs., 11 ozs., or 18,012 gm. Mr. Warren M. Foote of Philadelphia

very kindly sent the specimen to me for description and gave me the

foregoing information regarding its discovery and history.

The iron, which is a holosiderite, has been named Kingston from the

post office nearest to the place of discovery. As found, it was lenticular

in shape, and its dimensions were 204x167x70 mm. No sign of the

original crust remains, and the “thumb marks’ have been much ob-

scured by oxidation. The exterior, however, presents several broad, shal-

low depressions, which are clearly shown in Plates XXXV and XXXVI,

but nothing to indicate which was the briistseite. Crevices along which

oxidation has taken place to a considerable degree penetrate the mass

along the plates and are particularly noticeable in Plate XXXV, fig. 1

and Plate XXXVI, fig. 3. A combination of these cleavage crevices per-

mitted the breaking out of the fragment that was first submitted to assay.

The surface is thickly indented with oxidation pits 1-3 mm. across and

shallow in proportion to their diameter.

The material furnished me for investigation consisted first of the

whole mass and then of the two end pieces and seven slices into which

1 Published by permission of the Director, American Museum of Natural History.

(335)

336 ANNALS NEW YORK ACADEMY OF SCIENCES

the mass was first cut by the Foote Co., two fragments for specific grav-

ity determination and three for chemical study. The last, aggregating

about 50 gm. in weight, were sent to Booth, Garrett & Blair of Phila-

delphia for analysis, with the following result:

Per cent

He Cby: difference)! ihc sekck-t si sernes pains Bn cee 92.376

ING oie ertbene osc a ea EGET OC a an 6.980

LO Mth ea eam n teal ote ete aad val ey aise eins mc. DED 0.505

Os) Rete p ere ner Pies Sere TCA ACIRE ny RUN te oo 0.018

SSO ER ney Cater Orie pnts kits aan iaitn eo neuen aI es can Ree 0.014

Peieitas se ioeiey satis ta hava icaenctirey eee See eee Us OE Uae SC 0.099

Siig iat Sct Sule curs Bees CUD Stes rh elostea Sata UD te Ce 0.008

The analysts report the apparent presence of a trace of some other

element, probably of the iron-platinum group of metals, but state that

they were unable to isolate and determine it in the amount of material

available and that it must be present in variable quantities.

The specific gravity as determined at the American Museum on a

fragment weighing about 12 gm. is 7.63. This is a low value, even when

the small percentage of Ni + Co present is taken into consideration.

End piece No. 1 contains the angular hole left by breaking out the

fragment for assay. As submitted to me, this piece was about 40 mm.

thick and weighed 1174 gm. The Widmanstiitten lines (Pl. XX XVII)

were traceable over the whole polished and etched surface, but they

were obscured by a flecky granulation extending likewise over the whole

surface. The flecks, whose appearance reminds one of the particles

making up a flocculent chemical precipitate, are irregular in outline and

are from 0.4 to 0.5 mm. across, seldom reaching the latter dimensions.

They have no particular orientation and are so strongly developed in

places as to suggest an approach to ataxitic structure. ‘This portion of

the mass closely resembles Tazewell in appearance. 'The etched surface

of slice No. 1 was about 13 mm. distant from that of the end piece. It

showed the flocculent granulation over about three fourths of its surface

(Pl. XXXVIII), the remainder being occupied by a subcentral oval

area about 90 x 40 mm. in size in which the kamacite was practically free

from granulation. In the succeeding slices, the oval clear area ex-

panded until in slice No. 6 there was no granulated area (Pl. XXXIX).

End piece No. 2, however, showed a bright fleck here and there. The

granulated portion of the mass, therefore, originally formed a cap over a

roughly cone-shaped development of iron that was practically free from

the flecks. |

Turning now to the other features of the meteorite we may say that

the Widmanstiatten lines are well developed, forming kamacite bands

HOVEY, THE KINGSTON, N. M., SIDERITE 3317

from 0.5 to 1.5 mm. broad, but the usual width is 0.75 mm., where they

are not doubled or trebled. Some were measured that were practically

continuous for a length of from 75 to 85 mm. Neumann lines are abun-

dant and distinct, though occasionally obscured by a minute network

of curved lines. Teenite is practically absent and plessite is extremely

subordinate in development. ‘Thin, short lines of schreibersite may be

seen here and there, some of which are associated with little nodules of

troilite and some with the bands of kamacite. About fifty small nodules

of troilite were noted, varying in size from 1 to 8 mm. in diameter.

Lawrencite exuded rather freely from the crevices in the slices during

the dampness of summer.

The iron is octahedral in structure, and the breadth of the lamelle

throw it into the medium octahedrites (OM) of Brezina’s classification.

A circle whose radius is 70 miles would pass through or close to the

places of origin of the following New Mexico siderites: Kingston, Luis

Lopez (Magdalena), Oscuro Mts., El Capitan and Sacramento Mts.

The irons, however, seem to be independent falls.

ha? sete oe eo

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ANNALS N. Y. ACAD. SCIENCES

JOLUME NNIT, Puatp XXNXV

rant

& l[eustea

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ANNALS N. Y. ACAD. SCIENCES VOLUME XNII, PLare XXXVII

etched surface showing 2

which the granu-

fata)

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salsye Baottods - Pets. ‘oat

ANNALS N. Y. ACAD. SCIENCES VOLUME XXII, PLhaty XXXVIII

shed and etched surface from which —

tural size.

ANNALS N. Y. ACAD. SCIENCES VOLUME XNII, PLaty XXNIN

PUBLICATIONS

Spe 7 OF THE

_NEW YORK ACADEMY OF SCIENCES

(Lyceum or Natura History, 1817-1876)

: 1) The Annals (octavo series), established in 1823, contain the

ientific contributions and reports of researches, together with the rec-

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A volume of the Annals coincides in general with the calendar. year

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All publications are sent free to Fellows and Active- Members. The

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fA Subscriptions and inquiries concerning current and back numbers of

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THE LIBRARIAN,

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care of ‘

American Museum of Natural History,

New York, N. Y.

Editor, Epaunp Or1s Hovey

fogre)

H RTER, ‘CONSTITUTION AND. MEMBER- —

SHIP AN 1912

OF THE

2 nw YORK”

| PUBLISHED BY THE ACADEMY

20 a, 1913

THE NEW YORK ACADEMY OF SCIENCES

(Lyceum or Naturat History, 1817-1876)

OFFICERS, 1912

President—Emerrson McMiuur, 40 Wall Street

Vice-Presidents—J. EDMUND WoopM4N, FREpDERIC A. Lucas”

_ CHARLES LANE Poor, R. S. WoopwortH

Corresponding Secretary—HENry H. Crampron, American Museum

Recording Secretary—EpmunpD Otis Hovey, American Museum

Treasurer—HEnNRY L. DoHERTY, 60 Wall Street

Inbrarian—RawtpPuH W. Tower, American Museum

Editor—EHKpMUND OTIS Hovey, American Museum

SECTION OF GEOLOGY AND MINERALOGY

Channa . HE. WoopM4n, N.Y. University .

Secretary—CuarLEs P. Berkey Columbia University

~SHCTION OF BIOLOGY

Chairman—FREpERIc A. Lucas, American Museum

Secretary—WiLL1AM K. Grecory, American Museum

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY

Chairman—CHar.es LANE Poor, Columbia University

Secretary—F. M. PEDERSEN, College of the City of New York —

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY

Chairman—R. 8S. WoopwortH, Columbia University

Secretary—F REDERIC LiyMAN WELLS, Columbia University

The sessions of the Academy are held on Monday evenings at 8:15.

_o’elock from October to May, inclusive, at the American poe we of

Natural History, 7 7th Street and Central Park, West.

[ANNALS NEw YorK ACADEMY OF SCIENCES, Vol. XXII, pp. 339-414.

20 April, 1913.]

RECORDS OF MEETINGS

OF THE

NEW YORK ACADEMY OF SCIENCES.

January to December, 1912.

By Epmunp Oris Hovey, Recording Secretary.

LECTURE.

3 JANUARY, 1912.

Edward E. Barnard : THE PLANET Mars.

BUSINESS MEETING,

8 JANUARY, 1912.

The Academy met at 8:23 Pp. M. at the American Museum of Natural

History, Vice-President Woodman presiding. °

The minutes of the last business meeting were read and approved.

The following candidates for membership in the Academy, recom-

mended by Council, were duly elected:

ActTIvE MEMBERSHIP.

A. B. Pacini, Board of Water Supply, New York.

Henry Arctowski, New York Public Library.

ASSOCIATE MEMBERSHIP.

Charles R. Fettke, Livingston Hall, Columbia University.

The Recording Secretary announced that a special fund had beer

provided so that during 1912 each of the four sections of the Academy

might have $100 at the disposal of the sectional officers for expenses or

honoraria connected with an effort to make the meetings more interesting

(339)

340 ANNALS NEW YORK ACADEMY OF SCIENCES

to the general public and extend the eee of the Academy. A

vote of thanks was extended to the unnamed donor.

The Academy then adjourned.

EpmMuNpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

8 JANUARY, 1912.

Section met at 8:28 Pp. m., Vice-President J. E. Woodman presiding.

Thirty-three members and visitors were present.

The minutes of the last meeting of the Section were read and

approved.

In the absence of Dr. Charles P. Berkey, Secretary of the Section,

Dr. E. O. Hovey was elected Secretary pro tem.

The Secretary pro tem presented an application in the name of Mr.

George Borup for a grant of $500 from the Esther Herrman Research

Fund, as a contribution to the Crocker Land Expedition which he and

Mr. Donald B. MacMillan were organizing under the auspices of the

American Museum of Natural History and the American Geographical

Society. On motion, the application was referred to the Committee

on Grants from Research Funds with power.

The programme for the evening was then taken up as follows:

A. B. Pacini, THz MeTAMoRPHISM OF PoRTLAND CEMENT. I.

Remarks were made by Professors Kemp, Woodman and Grabau.

The paper has been published as pages 161-224 of this volume.

Dr. EK. O. Hovey gave a brief summary account of the Washington

meeting of the Geological Society of America and a few of the paper

presented there.

Professor A. W. Grabau gave a similar account of the Washington

meeting of the Paleontological Society.

The Section then adjourned.

EpmMuND OTIs Hovey,

Secretary pro tem.

SECTION OF BIOLOGY.

15 January, 1912.

Section met at 8:15 Pp. m., Vice-President Frederic A. Lucas pre-

siding.

RECORDS OF MEETINGS 341

The minutes of the last meeting of the Section were read and

approved. ;

The following programme was then offered:

Henry Fairfield Osborn, PHYLOGENY AND ONTOGENY OF THE HoRNS

oF MAMMALS.

Henry Fairfield Osborn, SkuLL MrasurmpMENTS IN MAN AND TH

Hoorep MAamMMALs.

Frederic A. Lucas, WHALING IN THE OLDEN TIME.

SUMMARY OF PAPERS.

Professor Osborn said in abstract: The recent discovery of the modes

of origin of the horns in the titanotheres, a perissodactyl group remotely

related to horses, tapirs and rhinoceroses, permits of a comparison of

phylogenesis with the ontogenesis of the horns in bovine mammals.

The latter is based upon an osteological series recently prepared by Mr.

S. H. Chubb, the former is based on the rich phylogenic series of Hocene

titanotheres in the American Museum of Natural History. The conclu-

sion is that ontogeny closely recapitulates phylogeny, that the genesis

is gradual or continuous, that the horns arise definitely and deter-

minately. In the bovine series it seems, in accord with the conclusions

of Dist, that the horn first appears as a circular thickening of the skin,

accompanied by accelerated growth of the hair preparatory to the forma-

tion of the keratin of the horny substance, at a period considerably prior

to any sign of the horn in the bony structure of the frontals. This

raises the problem, which will form the subject of a special paper in

the Annals of the Academy, as to what element first arises in connection

with horn evolution, namely: (1) the psychic, or desire to use the horn;

(2) the epidermal callous or keratin protection of the bony horn center,

or (3) the bony or osseous horn itself. It would appear that the psychic

tendency must precede the epidermal and that the latter precedes the

osseous, but this disputed point requires further investigation.

The paper was illustrated with lantern slides, drawings and specimens.

Professor Osborn, in his second paper, said in abstract: Comparative

anatomists and zodlogists have been slow to introduce into mammalogy

systems of measurement by indices and ratios, which have proved of

such universal value in anthropology. It is found among the hoofed

mammals, from studies undertaken by the author with the co-operation

of Dr. W. K. Gregory, that cephalic indices and limb ratios between

different segments of the skeleton are far more significant than systems

of direct measurement. These cephalic indices of the gradual changes

of proportion between different regions of the skull have the value of

849 ANNALS NEW YORK ACADEMY OF SCIENCES

specific characters and sharply distinguish members of different phyla.

For example, in the cross between the horse (f. caballus) and the ass

(EZ. asinus), it is found that the cephalic indices are transmitted as pure

non-blending characters.

Among the most significant indices are the following: a) ¢ the cephalic,

which is obtained by dividing the total or basilar length of the skull

by the zygomatic breadth; (2) the cranial, which is obtained by divid-

ing the basilar length by the postorbital length of the skull; (3) the

facial, obtained by dividing the basilar length by the preorbital length

of the skull, ete. The horses show proopic dolichocephaly, or elongation

of the face, and a static condition of the cranium, while the titanotheres,

in contrast, show opisthopic dolichocephaly, or elongation of the cranium,

and abbreviation of the face. Like the phyletic differences of proportion

between the horse and the ass, these differences are most exactly ex-

pressed by the method of indices.

The application of the ratio method to the limbs of the hoofed mam-

mals has again produced most surprising results. It is found that mam-

mals of different phyla adapted either to “weight” or to “speed” con-

verge respectively toward typical “weight” or “speed” ratios, which are

obtained by dividing the length of the lower segments, tibia and radius,

respectively, by the upper segments, femur and humerus, metacarpus

and metatarsus, respectively. These “weight ratios” and “speed ratios”

are far more significant as regards function and phyletic change than

the actual or direct measurements.

Dr. Lucas exhibited lantern slides illustrating some interesting pic-

tures from old works on whaling and showing the methods practiced by

the early Japanese, European and American whalers.

The Section then adjourned. WILLIAM K. GREGORY,

Secretary.

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY..

22 JANUARY, 1912.

Section met at 8:15 Pp. m., Vice-President Campbell presiding.

The minutes of the last meeting were read and approved.

Dr. F. M. Pedersen of the College of the City of New York was then

elected Secretary of the Section for the year 1912.

The following programme was offered :

C. C. Trowbridge, Recent DiscoveriEs CONCERNING A CHEMICALLY

Active Moprrication or NITROGEN.

William Campbell, Some Notrs on Iron AND STEEL.

The Section then adjourned. F. M. PEDERSEN, Secretary.

RECORDS OF MEETINGS 343

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

29 JANUARY; 1912.

Section met in conjunction with the American Ethnological Society

at 8:15 p. M., Gen. James Grant Wilson presiding.

The minutes of the last meeting were read and approved.

The following programme was then offered:

Pliny E. Goddard, Notrs oN THE JICARILLA APACHE.

SUMMARY OF PAPER.

Dr. Goddard in his paper said: The Jicarilla Apache are, from the

point of view of material culture, a buffalo-hunting Plains people dwell-

ing in skin-covered tipis. Their social organization differs from that

of the Navaho and neighboring Pueblo tribes in lacking exogamous clans,

there being two geographical divisions with ceremonial and political,

but not marriage-regulating, functions. Among the ceremonies the

speaker mentioned an annual feast celebrated on the 15th of September

and probably connected with the corresponding celebration at Taos, the

conspicuous feature of both consisting in a relay race. A ceremony

resembling the Bear Dance of the Southern Ute is performed in cases

of illness and is characterized, among other things, by .sleight-of-hand

performances of masked dancers. The girls’ puberty celebration is very.

prominent; a distinctive feature of the Jicarilla form of this ceremony

seems to be the association of a young man with the adolescent girl.

Among the myths of the Jicarilla that of the twin heroes is prominent.

In the course of the discussion Dr. Goddard stated that he had been:

unable to discover myths definitely connecting the mythology of the

Jicarilla with that of their linguistic congeners in California and the

Far North. In reply to another query he expressed his belief that,

owing to the linguistic differentiation of the Apache, this tribe must

have occupied its southwestern habitat a considerable period before the

first historical notice of it. (

The Section then adjourned. F. Lyman WELLs, Secretary.

BUSINESS MEETING.

5 FEpruary, 1912.

The Academy met at 8:21 Pp. M. at the American Museum of Natural

History, Vice-President Woodman presiding.

The minutes of the last business meeting were read and approved.

344. ANNALS NEW YORK ACADEMY OF SCIENCES

The following candidates for active membership in the Academy, rec-

ommended by Council, were duly elected:

Charles HE. Sleight, Ramsay, New Jersey.

R. B. Earle, New York University.

On motion, the following minute was unanimously adopted and or-

dered to be engrossed and transmitted to the family of the late Mr.

Charles F. Cox:

The Academy suffers irreparable loss through the death, on 24 January, 1912,

of Mr. Charles Finney Cox. For thirty-six years an Active Member and

Fellow of the organization, his influence has been felt ffom the first in all

progressive movements of the Academy. He served the Academy diligently as

Curator, 1884, 1885; Councilor, 1891, 1892; Treasurer, 1893-1907; President,

1908, 1909. At the time of his death he was again acting as Treasurer. When

President he was active in the organization of the Academy’s Darwin Cen-

tennial celebration, and he delivered a masterly address on Darwin at the

close of each of his two years of incumbency.

Always the friend of investigation, he was one of the founders of the Scien-

tific Alliance of New York, the first association of the scattered organizations

that were striving independently to advance the interests of science in the city.

Some five years ago he was again active in establishing the closer affiliation

which now obtains among them.

Mr. Cox’s consuming interest outside of his daily duties in the railways of

the New York Central system was the study of the life and writings of Charles

Darwin. In its pursuit, he became a keen and devoted collector of Darwiniana,

and the portraits, first editions, manuscripts and other priceless memorials

which he brought together constitute a remarkably complete exhibit of Dar-

win’s scientific work and influence upon the thought of the last fifty years.

Another of his avocations was microscopy, in which he was active for many

years, while his interest in botany was evidenced by his participation in the

founding of the New York Botanical Garden and in its management up to the

time of his decease. ;

In character, Mr. Cox was a man of great simplicity and natural refinement.

He attracted. and held his friends with bonds of attachment that were

altogether exceptional in their strength. While he will be missed and mourned

by all who knew him, the sense of loss is peculiarly deep in the circle of the

New York Academy of Sciences.

To his family the Academy extends its profound sympathy.

The Academy then adjourned.

EpMunpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

5 FEBRUARY, 1912.

Section met at 8:31 Pp. M., Vice-President Woodman presiding.

RECORDS OF MEETINGS | 345.

In the absence of the Secretary, Dr. Hovey was elected Secretary

pro tem.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered:

E. O. Hovey, THE Amaia Farm METEORITE.

Vernon F. Marsters, A SKETCH OF THE PHYSIOGRAPHY AND EARLY

Mining DEVELOPMENTS OF PERU.

Henry 8. Washington, RELATIONS OF THE FELDSPARS, LENADS AND

VEOLITES. :

George H. Girty, On Some FossILs OF THE LyKINS FORMATION.

(Read by Title.)

SUMMARY OF PAPERS.

Dr. Hovey exhibited a polished and etched slice of the iron meteorite

from the Amalia Farm near Gibeon, Africa, and called attention to the

interesting curvature of the Widmanstatten lines in certain portions

of the slice, apparently due to the softening of the neighboring surface

of the mass as it passed through the air; also a line of discordance be-

tween the lamella apparently due to welding by impact of two masses or

two fragments of the same mass before the meteorite reached the earth.

Mr. Marsters in his paper described the coastal plains and cordilleras

of Peru and gave sections at several points from the sea to the summit of

the eastern range of the Cordilleras, the petroleum deposits along the

coast and the great deposits of coal, Lake Titicaca and the mines vf

gold, silver, copper and vanadium along the contacts of the eruptive

rocks with the sandstones and the shales of the middle and eastern

Cordilleras.

Dr. Washington in his paper gave an ingenious regrouping of the

molecules in the standard analyses of the feldspars and related minerals,

bringing out the isomorphism of the groups more clearly than is done

by other methods of writing the formulas, provided one can admit that

silicon acts as a base as well as an acid.

The Section then adjourned.

EpmMuND Otis Hovey,

Secretary pro tem.

346 ANNALS NEW YORK ACADEMY OF SCIENCES

SECTION OF BIOLOGY.

12 Fesruary, 1912.

‘Section met at 8:15 Pp. M., Professor Bashford Dean presiding.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered :

IN SoutTH AMERICA.

This paper has been published as pages 9-112 of this volume.

The paper, which was illustrated with maps and diagrams, was dis-

eussed by Professor Dean, Dr. W. D. Matthew and others.

The Section then adjourned.

WILLIAM K. GREGORY,

Secretary.

‘SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

26 FEBRUARY, 1912.

Section met at 8:15 P. M. in conjunction with the New York Branch

‘of the American Psychological Association, R. S. Woodworth serving as

‘chairman.

The minutes of the last meeting of the Section were read and

‘approved.

The following programme was then offered:

3. E. Hickman, THE INFLUENCE oF NARCOTICS ON PHYSICAL AND

MENTAL TRAITS OF OFFSPRING.

A. KE. Chrislip, AUDITORY AND VISUAL MEmory.

Henry H. Goddard, THe Herrepity or Mentat Tratrts.

SUMMARY OF PAPERS.

Dr. Hickman said in abstract: The purpose of the study was to learn

if the use’ of narcostimulants (tea, coffee, tobacco and alcohol) had any

‘effect on the offspring. The research extended over a period of four

-years. It included 306 families with 2,560 children; 620 of this num- —

‘ber were students of Murdoch Academy, Utah. These were carefully

‘measured by medical experts and teachers to get their physical and

imental status. The measurements and examinations included height,

RECORDS OF MEETINGS B47

weight, eyes, ears, nose, throat, teeth, heart, lungs, stomach, spleen,

liver, kidneys and nervous condition. A record of the death-rate in

the families was obtained as well as a record of the student’s intellectual

standing. The students were divided into eight claasses, according to

the kinds and quality of stimulants used by the parents.

The examination showed: first, that there was on an average a very

decided difference between the offspring of abstainers and those of users,

even where tea or coffee was used by only one parent, for the offspring

of the abstainers were superior in size, intellect and bodily condition to

those of the caffein parents; secondly, as the use of caffein was increased

by the parents, from once to three and four times a day, a gradual de-

erease in height, weight, bodily condition, etc., of the offspring was

manifest ; thirdly, in families where not only tea and coffee were used,

but also tobacco, the children were still more inferior mentally and

physically, increasingly so with the increase of caffein drinks in connec-

tion with tobacco; fourthly, where alcohol was used with the above

narcostimulants the lowering of the physical and mental status was very

marked.” :

Comparing all the offspring of the narcostimulant parents with those

from abstaining parents, the latter were found to be better in all the 22

measurements than the former. Some of the differences were very

great, especially in weight, height, eyes, ears, physical health and rate of

mortality. There are over 100 per cent more eye, ear and physical de-

fects in the offspring of narco-parents; 72 per cent more children died

in this than in the abstaining class; 79 per cent of the narcostimulant

families had lost one or more children, while only 49 per cent of the

abstaining class had lost any children. It was also shown that the

death-rate of the parents in this latter class was 41 per cent higher than

in the former. The research also brought out the fact that it took the

ofispring of the narcostimulant parent eight tenths of a year longer to

graduate from the grades. In the Academy they were on an average a

year and seven months older than the students from the abstaining class.

Mr. Chrislip said in abstract: Experiments have been carried on in

the psychological laboratory of Columbia University and elsewhere for

the purpose of comparing visual and auditory memory. The points in-

vestigated in the first experiment were to determine: the number of

repetitions required by each sense to reproduce in a certain order cer-

tain total series of like construction; the average number of characters

of a series recalled in their proper order for each repetition of series of

like construction for each sense; and to determine, if possible, the best

material for testing the two senses.

348 ANNALS NEW YORK ACADEMY OF SCIENCES

The material used consisted of numerals, nonsense syllables and

words. Series composed of 12 and 16 characters of each material were

used in testing both senses.

The result shows that when series of 12 numerals similarly con-

structed were presented to the two senses, that out of 26 cases 20 are

visual, 8 auditory and 8 show no difference. In the case of the series

of 16 numerals, 19 visual, 4 auditory and 13 show no difference. With

12 nonsense syllables there are 15 visual and 15 auditory, the rest show-

ing no difference, but for 16 nonsense syllables, 25 visual, 7 auditory

and 4 show no difference. With the 12 words there are 14 visual, 10

auditory and 12 no difference; with 16 numerals, 22 visual, 9 auditory

and 5 show no difference.

For each repetition of each series the result shows that in the mem-

ory tests for visual reproduction the greater average number is repro-

duced. The nonsense syllables were the best material, as they offered

few combinations or devices for memorizing them.

Experiments, in which stories of 100 words each have been used to

test the two senses, have been carried on for some time. The two senses

have been tested for both immediate and delayed recall. In both the

immediate and the delayed reproductions the visual has been better than

the auditory. There is an experiment now in operation in which the

method is somewhat different from that in the former experiments con-

ducted with logical material. While the results are not all deter-

mined the indications are that the auditory may surpass the visual.

Dr. Goddard said in abstract: It is not the purpose at the present

time to present any results, but rather to make some suggestions and

point out possible lines of research in the hereditary transmission of

mental traits which may be of interest to psychologists.

In connection with our studies of the cause of mental deficiency at the

training school at Vineland, much material has been accumulated show-

ing the hereditary transmission of deficiency. In connection with these

data many facts have come to hand which make it clear that not only

deficiency, but many positive traits are directly transmitted. It is fur-

ther suggested that psychology would gain valuable data and contribu-

tions to many of its problems from a study of this question of heredity.

Indeed, it seems quite possible that many problems which are now so

complex as to elude our powers of analogy would be easily analyzed if

we were able to study the heredity problem and thus eliminate the

hereditary factor. For example, if the goodness of memory depends, as

Professor James said, upon the natural retentiveness of the brain tissue

plus the logical association that the individual establishes, then we may

RECORDS OF MEETINGS 049

reasonably expect that the condition of the brain tissue may be a quality

that is transmitted and could be eliminated through the study of mode

of transmission ; or, in other words, we could determine to what extent

the differences in memory are due to acquired factors.

It would seem equally possible that sensory conditions may be traced

through families, just as peculiar eyes or eyesight, peculiar hearing,

kinesthetic sensations, taste, or smell may be dependent upon organic

conditions which may be found to be directly transmitted. The inborn

habits or instincts are so bound up with acquired habits that it makes a

very complex problem. It seems quite possible that a study of the in-

stinctive activities of members of different generations might reveal to

us a good deal about the nature of instinct and its transmission which

would have very important bearings upon many of our problems of in-

stinct and emotions. Even the study of such a. complex problem as the

inheritance of mental deficiency may possibly yield us some most im-

portant results. .

It seems hardly likely that mental deficiency is due to the absence of

any one characteristic, but of several, and that it may be pictured more

as though normal mentality is the result of a hundred factors of which

a person must have, say, seventy-five in order to have what is called nor-

mal mentality. Now the twenty-five that are lacking may be any

twenty-five, perhaps, in the whole list and a tracing of the hereditary

traits might lead us eventually to determine some things about the re-

sulting mentality when the missing factors belong to different groups.

We shall work on these problems at Vineland as rapidly as possible,

but they should be studied in normal people as well. It is perhaps true

that it would not be possible to go back farther than the living genera-

tions ; but even so, if careful studies and tests were made of the mental

traits in living persons, it would be possible to get the records of two

and sometimes three generations, and these records could then be kept

and supplemented as the years go by and the newer generations come

on. There would thus be laid the basis for most valuable studies

later on.

The family histories, that we have secured in connection with our

children at Vineland, suggest two or three interesting questions. For

instance, there are several families in which alcoholism is strong in sev-

eral generations. It is possible that we have in these families an un-

usual appetite for alcohol, which appetite has been transmitted. It

looks as though it would not be impossible to eliminate to quite an ex-

tent the environmental factor, and so be able to determine whether this

was hereditary or not. The same is true of the sexual life. A great

a ae

oar

350 ANNALS NEW YORK ACADEMY OF SCIENCES

many charts show very much sexual immorality: and possibly here we

may have, in some cases at least, an unusual development of the sex in-

stinct which has broken over all bounds of conventionality and has

shown in different generations. It appears that all of these problems

are not only worthy of study, but might yield most important results.

The speaker showed graphic charts illustrating the family histories of a

number of families. ‘These charts showed the strong inheritance of

feeble-mindedness and also illustrated the points made in regard to

alcohol and sexuality. Considerable discussion followed.

The Section then adjourned. f

F. Lyman WELLS,

Secretary.

BUSINESS MEETING.

4 Marcu, 1912.

The Academy met at 8:16 p. m. at the American Museum of Natural

History, Vice-President Woodman presiding.

The minutes of the last business meeting were read and approved.

The Recording Secretary reported that Mr. Henry L. Doherty had

been elected Treasurer to fill the pnexpired term of Mr. Charles F. Cox,

- deceased.

The Academy then adjourned.

EpmunpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

4 Mancn, 1912. |

The Section met at 8:22 Pp. M., Vice-President Woodman presiding.

Fifty members and visitors were present.

The minutes of the last meeting a the Section were read, corrected

and approved.

The following programme was then offered:

J. E. Woodman, ForELANDS oF THE Bras p’OR LAKkEs, CAPE BRE-

TON IstAND, Nova Scorta.

Charles P. Berkey, Is THERE FauLT CoNTROL oF THE Hupson RIVER

CouURSE?

V. F. Marsters, DiIsTRIBUTION OF PETROLEUM DeEPosITs IN PERU.

SUMMARY OF PAPERS.

Professor Woodman illustrated his paper with many photographs

showing exceptionally fine views of the topography of the region and

the cliffs, bars and spits of the shores. 'T’wo peneplain surfaces are to

be seen—the higher, characterized by crystalline rocks, and the lower

by Carboniferous strata. The hooked spits and loops and bars of re-

_ cent wave and current work are developed on a scale that is seldom

equaled.

Professor Berkey discussed the reasons for the usual belief that the

straight course of the river is due to fault structure and illustrated this

view by the use of Professor Hobbs’s map of Atlantic border hneaments.

Tt was then shown that a detailed study of the structural geology of the

lower Hudson region shows many strong faults crossing the river

obliquely N. HE. to 8S. W. and some much less important running N. W.

and S. E., but none of any real consequence running N. and SB. or par-

allel to the Hudson River. It would be considered most éxtraordinary

if a great river like the Hudson should be controlled by small, insignifi-

cant faults and pay so little attention to the large fault zones.

Remarks were made by Professor Woodman. .

Mr. Marsters showed maps and drew sketches of structure and topog-

raphy. ‘Two belts of oil-bearing strata were described, both Tertiary—

a coastal region and an interior region.

The Section then adjourned.

CHARLES P. BERKEY,

Secretary.

SECTION OF BIOLOGY.

11 Maron, 1912.

By invitation of Professor C.-E. A. Winslow and his colleagues, the

Section met at 8:15 P. m. in the Department of Zodlogy of the College

of the City of New York, Vice-President Lucas presiding.

The minutes of the last meeting of the Section: were read and ap-

proved. —

The following programme was then offered :

C.-E. A. Winslow and I. S. Kligler, THz Number anp Kinps oF Bac-

TERIA IN Crry Dust.

C. V. Chapin, THe ARIAL TRANSMISSION OF

DISEASE,

852 ANNALS NEW YORK ACADEMY OF SCIENCES

SUMMARY OF PAPERS.

C.-E. A. Winslow and I. 8. Kligler in their paper presented the re-

sults of the examination of about 170 samples of dust from streets,

schools, houses and public buildings in New York. The total numbers

of bacteria found varied from 150,000 per gram to 145,000,000, aver-

aging from 3,000,000 to 5,000,000 from the indoor dusts and 49,000,000

from the street dust. Spores made up usually less than one-tenth of

the total. The count obtained at body temperature was about one-half

that at room temperature, averaging from 2,000,000 to 3,000,000 per

gram in the indoor dusts and 22,000,000 in the street dusts. B. coli

was usually present; in the street dust an average of 51,000 per gram

was found and in two samples over 100,000, while none showed less

than 100. The indoor dust, on the other hand, showed an average of

between 1,000 and 2,000. Acid-forming streptococci, such as are char-

acteristic of the mouth, were present to the extent of over 1,000 per

gram in three-fourths of the street samples and one-half of the indoor

samples. The average for the street samples was about 40,000 per

gram; for the indoor samples about 20,000 per gram. The large pro-

portion of these organisms, particularly in the indoor dusts, appears to

be significant of buccal pollution.

The paper, which was illustrated by charts and diagrams, was dis-

cussed by Dr. Lucas.

Dr. Chapin said in abstract: The diffusion of contagion through the

room or out-of-doors only was considered, not droplet infection, which

does not take place beyond a meter. Bacteriological evidence was not

discussed, though the quantitative work of Winslow on sewer air and

spray infection was referred to, a work which he is now extending to

dust. Epidemiological study and experiment have been rapidly nar-

rowing the list of alleged air-borne diseases. We now know that yellow

fever and malaria are never air-borne; experiments have shown that

bubonic plague and Mediterranean fever are not. There is no evidence

that cholera and typhoid fever are ever air-borne and much that they

are not. The spread of influenza out-of-doors does not take place, and

perhaps not indoors. The alleged evidence that smallpox virus is air-

borne around hospitals is very weak. Careful observation in hospitals

has shown that typhus fever, cerebro-spinal meningitis and poliomye-

letis do not pass from patient to patient in the same ward. ‘The same

is true for uncomplicated scarlet fever and for diphtheria except by con-

tact or close droplet infection. Probably measles and whooping cough,

rubella, mumps, chickenpox and smallpox are not air-borne, even in the

RECORDS OF MEETINGS 353

same room, but further observation may show that such infection may

rarely take place.

The paper was discussed by Professors Winslow and Bristol.

The Section then adjourned, the members visiting and examining the

laboratories and lecture rooms of the Department of Zoology, College

of the City of New York.

Witiiam K. Gregory,

Secretary.

LECTURE.

15 Marou, 1912.

F. S. Archenhold: Astronomy, EpucaTION AND CULTURE.

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY.

18 Marcu, 1912.

Section met at 8:15 Pp. M., Vice-President Poor presiding.

The minutes of the last meeting of the Section were read and ap-

proved.

The following programme was then offered:

Louis H. Friedburg, Propucrs or CHEMICAL ART.

SUMMARY OF PAPER.

Dr. Friedburg said in abstract: There are three lines along which

synthetic chemistry is to-day advancing. First, the production of

things found in nature; for example—wintergreen oil, vaniline and

camphor. Second, ennobling one substance into another; for exam-

ple—transformation of cellulose into artificial silk. Third, prepara-

tion of substances which are similar to natural substances, but which

are not found in nature; for example—celluloid and bakelite.

There are some important discoveries which have been made by acci-

dent, such as that of glass 2,600 years ago. The chemist must be alert —

enough to recognize the value of such accidental discoveries.

The speaker described in a very interesting and entertaining manner

the manufacture of parchment paper, mercerized cotton, gun cotton,

collodion and artificial silk. He showed on the screen many beautiful

examples of the Lumiére colored photographs, and the glass caterpillar

or spinneret which is used for making the artificial silk fibres. Reau-

mur in 1784 first suggested the possibility of artificial silk. Celluloid

bo4+ ANNALS NEW YORK ACADEMY OF SCIENCES

-and «heckerboard screens for photo-color printing, and, lastly, the

making of bakelite or artificial amber (so-called) were explained. Many

fine specimens of all substances mentioned were shown.

The Section then adjourned. ‘

F. M. PEDERSEN,

Secretary.

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

25 Marouw, 1912.

Section met in conjunction with the American Ethnological Society

at 8:15 Pp. m., General James Grant Wilson presiding.

The minutes of the last meeting of the Section were read and ap-

proved.

The following programme was then offered :

Robert H. Lowie, Dr. RapbosavLJEVIcH’s CRITIQUE OF PROFESSOR —

Boas.

SUMMARY OF PAPER.

Dr. Lowie stated that Dr. Radosavljevich had misrepresented Professor

Boas on a number of important points. He had entirely misrepresented

page 32 of Professor Boas’s preliminary report on “Changes in Bodily

Form of Descendants of Immigrants.”

The “economic” theory scoffed at by Dr. Radosavljevich is a figment

of his imagination. What Professor Boas says is that the arrivals dur-

ing the period following the financial panic of 1893 were under-devel-

oped in every direction. When Dr. Radosavljevich reproaches Professor

Boas for not studying the effect of American soil and financial panics.

on the same individuals during a period of time representing the age of

his subjects, he shows that he has not the faintest notion of what Boas‘is —

discussing in connection with financial panics.

Secondly, Radosavljevich’s contention that Boas’s own observations do

not support his theory of a change of type is a most naive instance of

conceptual realism. The conventional classificatory divisions of head

forms have for Radosavljevich an absolute biological value, and unless

the head forms of descendants of immigrants fall outside the conven-

tional class of their parents he refuses to admit a change in head form.

_ Dr. Lowie explained what statisticians and biometricians actually mean

by a real difference between two series.

In the discussion which followed, several visitors asked for informa-

RECORDS OF MEETINGS 355

tion relating to the nature of Professor Boas’s methods, which were ex-

plained by Dr. Goldenweiser, who had taken part in the investigation.

The Section then adjourned.

F. Lyman WELLS,

Secretary.

BUSINESS MEETING.

1 Aprin, 1912.

The Academy met at 8:25 Pp. M. at the American Museum of Natural

History, Vice-President Woodman presiding.

The minutes of the last business meeting were read and approved.

The following candidates for membership in the Academy: recom-

mended by Council, were duly elected:

ACTIVE MEMBERSHIP.

Mrs. Henry W. Hardon, 315 West 71st Street.

ASSOCIATE MEMBERSHIP.

F. F. Hahn, Columbia University.

The Recording Secretary then reported the following deaths:

Edward Russ, an Active Member for 5 years.

John B. Smith, an Active Member for 5 years.

A letter was read from Dr. Hermann Credner thanking the Academy

for the honor bestowed upon him by his election to Honorary Member-

ship in the Academy; also an invitation from W. W. Gilchrist, Jr.,

artist, to the friends of the late Mr. Charles Cox, to view a portrait of

the former Treasurer of the Academy at the Folsom Galleries.

The Academy then adjourned.

EpmunpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

1 Apri, 1912. :

Section met at 8:25 p. M., Vice-President Woodman presiding.

The minutes of the last meeting of the Section were read and ap-

proved.

The Secretary announced that Mr. Alfred H. Brooks, of the United

States Geological Survey, had been secured for the May meeting, at

356 ANNALS NEW YORK ACADEMY OF SCIENCES

which time he would give a public lecture on the “Geology and Mineral

Resources of Alaska.”

Dr. Hovey announced that the.seismograph which had been presented

to the Academy by Mr. Emerson McMillin had reached New York and

had been passed by.the Custom House.

The following programme was then offered:

Wallace Goold Levison, IntusTRATIONS OF MINERAL ASSOCIATIONS

BY MrAns oF CoLoR PLATE AND OTHER

PHOTOGRAPHS OF OPAQUE SPECIMENS.

A. B. Pacini, THE METAMORPHISM OF PORTLAND CEMENT.

dale

Charles T. Kirk, ALTERATIONS IN THE SNAKE River BASALTs.

(Read by title.)

SUMMARY OF PAPERS.

Mr. Levison said in abstract: On a previous occasion the writer pre-

sented to the Academy a “Note on Photographs of Minerals for illus-

trating Books, Papers and Lectures” (Annals N. Y. Acad. Sciences,

Vol. XII, pp. 661 and 663). The examples consisted of lantern slides

and prints of light-colored mineral specimens of cabinet size mounted

on standard size blocks with standard size labels and of microscopic

mounts in Rakestraws. (See “Report of Committee on Standard Sizes,”

N. Y. Acad. of Sciences, 1894.) The writer found it difficult to obtain

satisfactory photographs of highly colored minerals in or on colored

matrices, as such associations usually afford poor contrasts on ordinary

plates.

To produce representations of colored. minerals at that time hand-

painted or colored lantern slides or prints were the chief expedient.

Some time later the M. A. Seed Dry Plate Co. introduced its G. B.

P. R. (green, brown, purple and red) plates, which served better than

ordinary plates for lantern slides of certain colored minerals. Thus

malachite and prehnite looked well on the green, native copper on the

red and rusty or yellow-colored minerals such as the stilbite and cal-

cite from Upper Mt. Clair, N. J., on the brown plates. The entire plate

was usually of a tint much similar to that of the specimen, but different

parts of the specimen usually developed in tints intermediate between

the four possible colors above mentioned, so that pictures on these ‘plates

made desirable lantern slides. This method was not applicable to paper

prints.

The recently introduced color plates of Lumiére, Jougla and Dufay

RECORDS OF MEETINGS SOK

now afford remarkably satisfactory lantern slides of colored minerals

of either cabinet or microscopic size.

The methods of production of all these plates were explained, but in

the writer’s experience the Jougla and Dufay plates seemed to afford

slides preferable in transparency and resistance to the heat of the

lantern.

Photomicrographs in color of thin sections of rocks and minerals by

transmitted polarized light were made by Francois Frank on Lumiére

plates as early as the year 1907 (Ch. A. Francois Frank, “La Micro-

photographie en couleur avec les plaques autocrome de M. M. A. et

L.. Lumiére,’ Comptes Rendus 1** Semestre, T. CXLIV, No. 24, p.

134L, 17 June, 1907).

The first attempt to make photomicrographs, on autocolor plates, of

microscopic colored minerals by ordinary reflected light was made by

Mr. Frank La Manna, of the Borough of Brooklyn, N. Y., about Feb-

ruary, 1911. Mr. La Manna thus photographed on Jougla and Dufay

plates he brought from Paris several specimens of microscopic colored

minerals mounted in Rakestraws by the writer. (F. La Manna, Ex-

hibit at the Annual Reception of the Department of Microscopy of the

Brooklyn Institute, 11 March, 1911.) The deep black interior of the

Rakestraws served as a superior black background.

Through the courtesy of Mr. La Manna, the writer received some of

these plates upon which he photographed other similar specimens.

These photomicrographs, jointly with those of Mr. La Manna, were ex-

hibited before the New York Mineralogical Club in April, 1911, and

again at the reception of the Brooklyn Institute’s Department of Micros-

copy, March 9, 1912, and with additions in illustration of this paper. ~

(W. G. Levison, Exhibit at the Annual Reception of the Department

of Microscopy of the Brooklyn Institute, March 9, 1912.)

In making these photomicrographs the writer used a lens with a small

stop giving a desirable depth of focus, a long bellows, a suitable color

screen, long exposure, reflecting screens to soften the shadows and a

very rigid adjustment of the apparatus and its supports.

These autocolor plates have likewise afforded the-writer very satis-

factory lantern slides of colored cabinet specimens. Each picture ob-

tained is a direct positive. Such positives may afford approximately

similar copies by the camera or other color plates, but duplicates made

directly from the specimen are preferable. They may also, like any

colored transparencies, be copied by contact on approximately similar

colors, on a paper called T'to color paper by Dr. J. H. Smith, recently

introduced from Paris.

358 ANNALS NEW YORK ACADEMY OF SCIENCES

Mr. Pacini’s paper has been published as pages 161-224 of this volume.

Remarks were made by Mr. Johnson and Mr. Price and Mr. Gaines

of the Board of Water Supply testing laboratory. Several questions

were asked by Professor Arnold of New York University. Remarks

were also made by Professor Arnold.

The Section then adjourned.

CHARLES P. BERKEY,

Secretary.

SECTION OF BIOLOGY.

8 APRIL, 1912.

Section met at 8:15 Pp. m., Vice-President Lucas presiding.

The minutes of the last meeting of the Section were read and

approved. .

The following programme was then offered:

Thomas H. Morgan, Sex-Linkep INHERITANCE IN POULTRY.

Louis Hussakof, THE Spawnine Hasits oF THE SEA LAMPREY,

Petromyzon marinus. i

John T. Nichols, Notes oN CuBAN MARINE FISHES.

SUMMARY OF PAPERS.

Professor Morgan’s paper has been published as pages 113-133 of this

volume.

Dr. Hussakof said in abstract: The observations were made on the

Nissequoque River at Smithtown, Long Island, June 1 and 2, 1911,

while collecting material for an exhibition group of Petromyzon for

the American Museum. The nests are depressions in the gravel of the

river bottom, two or three feet in diameter, and six inches deep at the

center. ‘The method of their construction and the general behavior

of the specimens of the nest are very similar to those of the Brook

Lamprey. But, owing to the large size of this species, all its movements

can be minutely observed.

The speaker exhibited a small model of the Lamprey group now under

construction in the American Museum of Natural History and also life-

size models of adult lampreys. The paper was also illustrated by lantern

slides.

Mr. Nichols dealt with the results of a brief collecting trip to Cuba

and exhibited various specimens. He passed in review some of the

Scombriform fishes. The king fish, Scomberomorus cavalla, is highly

esteemed, but another species, S. regalis, is said to be occasionally

RECORDS OF MEETINGS 359

poisonous. S. maculatus, the Spanish mackerel, was not seen. While

regalis and maculatus occupy more or less distinct areas, cavalla is

abundant both in Florida, with maculatus, and in Cuba, with regalis;

in the speaker’s opinion these two last-named species, which are still

closely related, have recently become separated through the competition

on cavalla. Two very widely separated forms, Arbaciosa rupestris and

Gobius soporator, were found inhabiting adjacent rock pools; both were

concealingly colored and could have been confused until their distinctive

color patterns were noticed.

The paper was illustrated by means of lantern slides.

The Section then adjourned.

WILLIAM K. GREGORY,

Secretary.

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

22 APRIL, 1912.

Section met at 8:15 P. M., in conjunction with the New York Branch

of the American Psychological Association, R. S. Woodworth acting as

Chairman. The afternoon session was held at the Psychological Labo-

ratory, Columbia University, and the evening session was held at the

American Museum of Natural History.

The following programme was offered :

Gertrude M. Kuper, InpivinuaL DIFFERENCES IN THE INTERESTS OF

CHILDREN.

T. H. Kirby, PRACTICE IN THE CASE OF CHILDREN OF SCHOOL

AGE.

C. D. Mead, THE AGE OF WALKING AND TALKING IN RELATION

TO GENERAL PRACTICE.

G. C. Myers, SEX DIFFERENCES IN INCIDENTAL Memory.

A. J. Culler, RELATION OF -INTERFERENCE TO ADAPTABILITY.

E. 8S. Reynolds,

J. T. Gyger and

L. L. Winslow, EXPERIMENT IN THE CaTcCHING or PENNIES.

D. O. Lyon, THE OPTIMAL DISTRIBUTION OF TIME AND THE

RELATION OF LENGTH OF MATERIAL TO TIME

TAKEN FOR LEARNING.

SUMMARY OF PAPERS.

Miss Kuper said in abstract: That interest plays a very important

dynamic role in the educational field is only too evident from such

360 ANNALS NEW YORK ACADEMY OF SCIENCES

treatises as Dr. Dewey’s article, “Interest as Related to Will” and Dr.

Montessori’s “Pedagogia Scientifica.” But interest is a general term

and can not have an absolutely universal value for every individual or

every subject of thought or desire. Individual interests are as important

in the social world as are individual capacities. They should, therefore,

be a fruitful field for scientific investigation. 'The experimental work

done with advertisements has brought to light group differences in the

preferences of men and women for various appeals. The investigation

to be reported was of a like nature, except that it dealt with children.

The formal experiment consisted in asking an individual child to

arrange nine pictures in the order in which he liked them best. The

nine pictures were chosen to represent nine specific appeals: landscape,

children, animals, religion, pathos, sentiment, patriotism, heroism, and

action. (They were Cosmos prints and therefore of uniform size and

finish.) In all, there were three series of these pictures, each parallel

so far as possible with the other two in their appeals. The children

numbered over 200, 10 girls and 10 boys for each year’s age from 6.5

to 16.5. They were almost entirely attendants of the public schools of

New. York City and came from quite varied sections of the city.

The results were tabulated according to age differences, broad social

distinctions, and nationality. In the last-named case the number of

subjects was so limited (10 girls and 10 boys to each of the following

nationalities: Irish, French, German, and Italian, and only 9 girls and 8

boys to the Spanish) that the results are not held as significant.

The positive data showed a sex difference in the order of preference

for these several appeals. The girls’ order was: (1) Religion, (2)

patriotism, (3) children, (4) pathos, (5) animals, (6) sentiment, (7)

landscape, (8) the heroic, (9) action. The last two were decidedly lowest

in the scale and the first three were quite clearly highest for all ages;

but the picture representing these nine curves was one of bewildering

intersections as the values changed from year to year. The boys’ order

was: (1) Religion, (2) patriotism, (3) action, (4) the heroic, (5)

pathos, (6) animals, (7) sentiment, (8) landscape, (9) children. The

boys’ chart representing the curves for these appeals showed greater —

agreement from year to year. Religion and patriotism, the heroic and

action, and landscape and children kept rather parallel courses all along

the age scale, and no very decided tendencies appeared with progressive

age differences. Girls seemed to lose interest somewhat in pictures of

children and animals and to take greater interest in the heroic and action

pictures. The latter change is explained by the fact that, as the girls

increased in school knowledge, they read an historical background into

these more or less warlike scenes.

RECORDS OF MEETINGS 361

A great sex difference was found in the variability measures, as cal-

culated for the various ages, appeals, social classes, and nationalities.

In every case but two, the girls exceeded the boys in their P.E.; and

in these two exceptions the boys’ P.H. was once greater than the girls’

by only 5 per cent., and another time exactly equal to the girls’ P.H.

The amount of sex difference was, as a rule, anywhere between 12 per

eent. and 57 per cent. This held true in every scale, whether according

to age, appeals, social class, or nationality. The girls’ average P.E. was

1.66; that for the boys was 1.36.

Both girls and boys were least variable about the subjects they liked

best, 7. e., religion and patriotism; but apart from these appeals there

was no correlation of variability with relative likes or dislikes.

It is a noteworthy fact that in range of variability the boys far ex-

ceeded the girls. The limits for the boys’ P.E. were .82 (patriotism)

and 1.60 (landscape), giving a range of difference of 78 per cent.; the

limits for the girls were 1.47 (religion) and 1.95 (animals), showing

a range of only 48 per cent. In this particular experiment this in-

dicates that boys are very much more agreed about some likes than are

girls, and yet quite as varied about others. In other experiments such a

range of variability may point to greater individuality of the male sex

among themselves while as a group they are relatively homogeneous.

Another sex difference noted was the number of positive dislikes

expressed by each sex. The girls gave 161, or 6 per cent. dislikes as

against the boys’ 65, or 2.4 per cent. Boys seemed to entertain relative

indifference toward the appeals at the bottom of the list. The things

the girls disliked most were (1) scenes of action suggesting death and

(2) pictures showing angry attitudes. The reasons given by the boys

for their dislikes were (1) gloomy, indistinct scenes, (2) sentimental

pictures, (3) costumes worn by men which were feminine in style or

left the figure partly nude, and (4) pictures suggesting illness.

A certain age difference revealed itself in the remarks made by the

children about the pictures. The seven and eight year olds showed

limited powers of observation. Some detail, and, in landscape scenes, al-

ways the human detail, no matter how small, was made the focus of

attention to the complete overlooking of the larger subject. Unfa-

miliar details when pointed out to them received as many different in-

terpretations as there were children. As the children grew older their

remarks of both girls and boys. Emotional attitudes, actions and even

of the pictures and they drew upon all their known sources for filling

in their perceptions. At the ages between 11 and 13 the critical spirit

made its first appearance among the girls. Only at fourteen did it

oceur in the boys’ comments. At these ages the emotions prompted the

362 ANNALS NEW YORK ACADEMY OF SCIENCES

remarks of both girls and boys. Emotional attitudes, actions and even

words were ascribed to the pictorial persons. At 15, the remarks be-

came more laconic, but what was said was significant and definite as

to the persons, place and action of the picture. This age marked the

first signs of hesitation in speaking of the pictures of sentiment. Up

to the age of nine the remarks had been very naive; after that the

pictures were dismissed with the phrase, “they’re lovers” or “a love pic-

ture”; often the characters were named Romeo and Juliet, Paul and

Virginia, etc.

In all their comments the girls were far more personal than the boys.

The personal pronoun and references to their individual experiences

were the usual preface to their statements. With the boys it was quite.

otherwise; they discussed the picture as an objective thing, independent

of their conscious existence. Boys tended to locate scenes in definite

historical time and specific geographic places.

The effect of uncertainty about a picture, crudely averaged, was a

displacement of about five places toward the lower end of the scale.

Dr. Kirby said in abstract: This experiment was conducted to get

some information concerning (1) the value of the practise experiment

as a method for school work and (2) the value of practise periods of

different lengths.

339 fourth year children belonging to 10 different classes took part

in the practise, which consisted of adding columns, each of 10 numbers,

0’s and 1’s not included, as rapidly as was consistent with accuracy,

each child competing with his own past record. Seven different sheets

of columns of equal difficulty were used (Thorndike’s Addition Sheets).

In every case there was one hour of practise, but for different classes

this hour. was broken into 2214-, 15-, and 6-minute periods, an initial

15-minute period and a final 15-minute period being given to form

the basis for determining the gain per cent.

The hour’s practise for the 339 children taken as one group resulted

in an average gain of 55 per cent.; median gain of 48 per cent. In a

similar test with 19 university students, Professor Thorndike found

an average gain of 29 per cent., median 33 per cent. from about 53

minutes of practise, and said: “The amount of improvement in this

experiment may also add to our confidence that the method of the

‘practise experiments wherein one works at one’s limit and competes

with one’s past record may well be made a regular feature in many

school drills. Even if the same length of time produced in children a

percentile improvement, only half as great as here, the gain would still

probably be far greater than the gain by any of the customary forms of

drill.”

RECORDS OF MEETINGS 363

For the classes which took the hour’s practise in 2214-minute periods,

there was an average gain of 61 per cent., median 49 per cent.; in 15-

minute periods, average gain 55 per cent., median 43 per cent.; in

6-minute periods, average gain 54 per cent., median 44 per cent.

Dr. Mead said in abstract: 1. Data.—50 “normal” children (25

boys and 25 girls), averaging less than six years of age, of graduate

students of Teachers College and Columbia College. Ages were thrown

to the nearest month. Walking means: “To take a step unassisted.”

Talking means: “To use a word intelligently, 7. e., to associate the idea

with the object.”

Results——The median “normal” child begins to walk at 13.5 months,

with a probable error of 1.06 months. The chances are 999 to 1 that

the true median will not differ from the median obtained by more than

.66 month. The extreme range is from 11 to 30 months. .90 per cent.

of the cases fall between 11 and 17 months. The median “normal” child

begins to talk at 15.7 months, with a probable error of 2.83 months.

The chances are 999 to 1 that the true median will not differ from the

median obtained by more than 1.96 months. The extreme range is from

9 to 25 months. 90 per cent of the cases fall between 10 and 21 months,

with 18 months as the mode.

Il. Data.—145 “schoolable” children (boys and girls) of the Indiana

School for Feeble-minded Youth, in reply to the question on the personal

descriptive entrance blanks: “At what age did the child commence to

walk?” and 92 in reply to the question: “At what age did the child

commence to talk?”

_ Results—The median feeble-minded child begins to walk at 21.8

months, with a probable error of 7.56 months. ‘The chances are 999 to

1 that the true median will not differ from the median obtained by

more than 3 months. The extreme range is from 12 to 72 months. 90

per cent. of the cases fall between 13 and 50 months.

The median feeble-minded child begins to talk at 34.2 months, with a

probable error of 12.6 months. The chances are 999 to 1 that the true

median will not differ from the median obtained by more than 6.5

months. The extreme range is from 12 to 156 months (only one case

going above 108 months). 90 per cent. of the cases fall between 14 and

84 months.

Dr. Myers said in abstract: A test was desired wherein the thing to

be remembered should be merely incidental and where the focus of the

subject’s attention should be directed away from the facts to be called

for after the exposure of the stimuli, but where these facts would have to

enter, wholly or in part, into the experience of the subject. To this end a

864 ANNALS NEW YORK ACADEMY OF SCIENCES

list of six simple words were used as stimuli. The subject was told

that he would be given a spelling test and he was led to believe that it

would be a real test in speed and accuracy of spelling.

A practise test with digits was given for three successive times before

the real test began, to delude the subject as to the purpose of the ex-

periment. A dozen or more digits were pronounced at random so

rapidly that the subject could scarcely keep up in writing them. In

the midst of this series of digits the experimenter, without any warning,

gave the signal for the subject to turn the page upon which he was writ-

ing, and continued to pronounce digits at the same speed. The subject

was told that the words would be given in the same manner, but not quite

so rapidly. The following words were then pronounced: angel, pickle,

dirt, busy, onion, women. ‘The last word was pronounced in such a

manner that another word was expected by the subject, but the signal,

“turn,” was given instead, and the subject was told to write as many of

these words as he could remember, to place them in the order in which

they had been given, and to indicate by a line the place for each omitted

word. The time each individual required to reproduce the words was.

recorded by a stop-watch.

After testing over 100 individuals the writer applied the test to groups

of college, normal-school and public-school subjects. Aside from imme-

diate reproduction, records were secured after various intervals, ranging

from 14 hour to 3 months. In all such cases a practice test of rapid fold-

ing of papers was added. After the words were pronounced the papers

were promptly collected and the experimenter left the room. The sub-

jects thought the work was ended, but at various times the experimenter

reappeared and asked for the reproduction. The time for all group repro-

duction was limited to 1144 minutes..

The best results were secured immediately after presenting the stimuli.

Practically the same efficiency was shown for the reproduction after 6

hours as for that after 14 hour. But there was a decided fall after 7 days.

and a still greater fall after 3 months.

No appreciable difference was shown in efficiency between the lower

grades and the college students for immediate reproduction; but after:

various intervals there was a gradual decrease in efficiency with age.

Of the 1,515 subjects, 757 females and 758 males, only 29 of the

former and 18 of the latter reproduced the six words in exact order.

In all grades the females were markedly superior to the males, both

for the number of words remembered and for order. They had a higher

central tendency and were more variable than the males in the 5th, 6th,

7th, and 8th grades, while for the other groups the males were more:

variable.

RECORDS OF MEETINGS 365

108 other subjects were tested with 10 letters and digits. Here the girls

answered more, but the boys were better for order.

Mr. Culler said in abstract: The purpose of this experiment was two-

fold: to determine the effect of differently distributed practise series

upon learning given material; and to make observations upon the learn-

ing process in general.

_ The material to be learned was the path from the beginning to the

end of the Hampton Court maze. The paper (8 by 6 inches) on which

the maze was printed, was affixed to a board. Over it was placed a large

circular piece of cardboard, easily movable, having in the center a small

opening (5% to 11/16 inch) through which extended a pencil to mark

the course of the subject?s movement. At no time could the subject

see more of the maze than the part visible through the opening. At

the beginning of the experiment the subject was thus instructed: Pencil

is now at the entrance tothe maze; keep on moving until you reach

the end. Never cross a line; always keep to an open path. Mazes are

all the same and will be placed in the same position.

At each trial the time was recorded and number of errors was counted

and recorded. ‘To each subject were given 12 trials. Subjects were di-

vided into 6 groups as follows: 12 trials at one time, 6 on 2 successive

days, 4 on 3 days, 3 on 4 days, 2 on 6 days and 1 on 12 days. There

were 5 men in each group except the last, in which were 3. With regard

to time of day, subjects were divided into two groups: one group each

‘day for the required number of days, after lunch (1-2 P. m.): the

second group each day after dinner (7-8 p. M.). In comparing men of

the two groups no account was taken of this slight difference, as it was

considered practically negligible. Good light was uniformly provided.

The interval between successive trials of a subject at the same sitting

was 30-40 seconds.

Subjects were all graduate students, age frou 22 to 28.

Three classes of errors appeared: Wrong choice between alternative

courses, retracing when on right course, and (accidentally) crossing a

line. The first kind are major errors (value 1) and the other two kinds

minor (value 144). These are arbitrary values for computing results.

The major errors were counted as follows: There are 6 (or 7, depending

upon the course taken) places where choice must be made between

alternative paths of which only one is right. Each time the subject

moved from one of these places in a wrong path, 1%. e., away from the

goal, it was counted one error. Errors of retracing when on the right

path were usually small and due to defective attention or eyesight—

subject either thought he had accidentally passed an opening and moved

366 ANNALS NEW YORK ACADEMY OF SCIENCES

back to see, or on coming to a turn failed to notice the opening and

thought he had run into a blind alley.

The results are as follows:

I. TABLE OF ABSOLUTE TIME AND HRROR VALUES ATTAINED IN HacH GROUP

(The different groups are indicated thus: One—12, etc.; the word indicates

the number of trials each day, the figure the number of successive days. The

two columns show the average of number of seconds consumed and number of

errors made in the last three trials in each group; thus showing the relative

standing of groups at end of practice period. ‘The figures in parentheses show

relative position. )

Time, Errors,

‘ Per Cent. Per Cent.

Dy sree La NDR cre ORE RI Vata une N agian ts 50 (3) 4.8 (4)

WO == Gie iece) rte sot ccs vost mravo to nerwaa eles eieteinual Siero nett 61 (5) 5.2 (5)

TATOO ih evans iis se Gide sus Seco ea paoseie, eee eT 59 (4) 3.2 (8)

QU OU Re cog Maan tot UW AR hea ea aA 39 (1) 9 (1)

NSD. aaa tas Ant tye | EERE ae iran es aM oeates Ree oe ae 75 (6) 5.5 (6)

Ce Ly ahs os UCL gO Be Ala eA Oa 48 (2) 3.0 (2)

II. TABLE OF PERCENTAGE GAINS

(In each ease the percentage represents the ratio between the average of

first three trials and last three trials in the same group. This table is intended

to show improvement of each group irrespective of absolute values attained.)

Time, Errors,

Per Cent. Per Cent.

Or aaa eta ante eae bas ogeeer cence pene 210.0 (4) 147.9 (5)

ID O=—=G wis, ol spel cnsicets eleicus encuecersetsencisianctens 253.0 (3) 161.5 (4)

PAT C GA 5 aviaisira te a. 'or0f sila staves cle ete ce esiees 195.0 (6) 802.0 (1)

MOUTH Gh aed abs cyselereieiey sven ered ones 341.0 (2) 218.5 (3)

SS TiO icin Crete SU aestatnaeiere anette ra Noalisveviens 206.6 (5) 125.3 (6)

MW OLV Oa 0 sii 28, sole eof evedtseley see! svelte 368.7 (1) 236.6 (2)

(It must be said that the results of Six—2 were vitiated by the professed

indifference of one subject, because of which both time and errors for the last

few trials in that group are abnormally high.)

The results seem to point to the following conclusions: In general,

outside the Six—2 group, the One—12 and Two—6 groups made the

lowest absolute records and also least improvement; this apparently in-

dicates that the learning period was too prolonged, with insufficient

practise at any one time. On the other hand, the ''welve—1 and Four—

3 groups show in general the highest absolute records and greatest im-

provement. Here the practise was more thorough each time and not

so prolonged. ‘The curve of greatest regularity is the Four—3 curve.

The three groups, then, in which practise periods were longer and

confined to a few days show better results than the three in which

practise periods are shorter and prolonged over 4-12 days. The applica-

RECORDS OF MEETINGS 367

tion to learning any material would seem to be that better results are

secured by a few more prolonged or persistent periods of study repeated

perhaps for several days than shorter periods prolonged over a greater

number of days.

Some observations were made on individual methods of learning

which can not be included here.

Messrs. Reynolds, Gyger and Winslow. The authors said in abstract:

The experiment had two aims: (1) To investigate the learning process.

(2) To find what transfer, from the right hand to the left, if any,

would be shown.

Three subjects took part in the experiment which follows. It was

carried on in two series: (1) That in which the subjects caught the

pennies, two at a toss, palm of the hand down. (2) That in which

they caught three. The first series was of 7 days’ duration; the second,

10 days’. The time for tossing was from 1 P. M. to 2 P. M. on Mondays

and Wednesdays. Conditions were as nearly constant as possible, the

same room being used throughout the experiment. In the case of the

two-penny series, the subjects caught for 10 trials and then rested for 10.

In the three-penny series two subjects caught at the same time, the third

subject resting. In the first case, score was kept by the two unemployed

subjects in turn; in the second case, by the one unemployed subject.

Certain conditions influencing accuracy were noted, among which

are the following: Some parts of the room were more conducive to

accurate catching than others, that nearest the window being the most

favorable. The pennies could be caught with most accuracy if no ob-

jects were in front of the subject to distract his attention. The tossing,

when carried on before a blank, light-colored wall, was most successful.

An increase in confidence and in accuracy resulted when a window was

opened to admit new air. An interruption, as that caused by another

person entering the room, was followed by a corresponding fall in score.

The subject, by counting to himself his successful tosses, was stimulated

to a better score. The nervous feeling of haste as well as nervousness

caused by outside matters of importance to the subjects (such as press-

ure of work) tended rather to increase than to diminish their scores.

Each subject discovered and followed his own methods of tossing.

After finishing the two series, the subject who had followed the method

of throwing his pennies high into the air was able to catch an additional

penny (making four in all) with very little effort. The other subjects

tried this continually and failed, their hands striking the floor before

the fourth penny was reached. The quick shutting of the hand was an

important factor. One subject was materially helped by thinking of

368 ANNALS NEW YORK ACADEMY OF SCIENCES

the word “grab” previous to each trial. In some instances, the second

penny would be caught and lost, the first and third being retained.

Although occasionally a subject would catch all three successfully with-

out knowing it, yet the tossing can not be said to have become automatic.

The progress in learning was unsteady. Yet in each case there was

a gradual advance, noticeable particularly in the beginning. A warming--

up period was universally experienced by each subject at the beginning

of each day’s practise.

In the second series, a transfer test was tried with the left hand

before and after the practise series. This showed a considerable increase

in ability to catch with the left hand.

AMOUNT OF TRANSFER CATCHES

Subject. Before Test. After Test. Per Cent Gain.

x NaS cecoiese cape prcee eareauentts seuievradiay euetrameul ae taste 3 14 4662

Dive aietasatera aides So hclng Smee gaa eae ee 11 32 290+

SD ta la asl aves cheers sal ot antebel aanare tele che ia edaae arenes 1 29 2900

Total gain......... eenoeenno 15 és) 500

Mr. Sax said in abstract: Although art and science are widely sepa-

rated, they may co-operate in art education. Prevailing methods are in-

direct, depending upon a never certain transfer of training. During

the three years the average student spends at art school, his course is as

follows: Casts and still life in charcoal; still life in color; anatomy and

perspective as formal subjects; the figure in charcoal; some composition,

and, finally, painting the head and figure in oils. |

Results show little transfer; for example, compositions show little

knowledge of anatomy or perspective. Charcoal and oils have few

identical elements in substance or procedure; in. fact, specific habits

formed in mastering charcoal often act preclusively when the student

attempts to paint. Students who can draw, but not paint; construct,

but not compose, or are draughtsmen, but not colorists, and their oppo-

sites are in the overwhelming majority.

Experiments now under way on the learning process as applied to

painting seem to show that (a) preparation in charcoal and still life*is

unnecessary in painting figures; (b) efficiency depends largely upon

correct analysis; (¢) muscular coordination plays a minor part; (d) a

direct method and generalized idea of procedure are essential and (e)

the control of attitude is most important.

Dr. Lyon said in abstract: This paper was divided into two parts, it

being in reality a discussion of two distinct questions: (1) “The Distri-

bution of Time in Relation to Economy in Learning and Retention” ;

RECORDS OF MEETINGS 369

and (2) “The Relation of Length of Material to Time Taken for Learn-

ing.” Concerning the first of these, it was shown that in estimating

economy, not only must we consider the time spent, but the degree of

retention as well. It was shown that individuals differ greatly, and that

where one could learn a set of ten stanzas in less time by the continuous -

method (i. ¢., doing the work in “one sitting”), another individual

could lower his total time by dividing the time spent into several periods,

é. g., by spending 5 minutes per day. With but three exceptions re-

tentiveness was decidedly better by the divided-time-method. This was

notably the case with nonsense-syllables and poetry. The most general

statement that can be made, taking all materials and methods of presen-

tation into consideration, is that the most economical method is to dis-

tribute the readings over a rather lengthy period—the intervals between

the readings being in arithmetical proportion. For example, with one

individual in memorizing a poem of twenty stanzas the highest retentive-

ness was obtained by distributing the readings as follows: 2 hours, 8

hours, 1 day, 2 days, 8 days, 16 days, 32 days, ete. The practical bearing

of the results obtained on education in general was then considered. The

above individual found that the most economical method for keeping

material once memorized from disappearing was to review the material

whenever it started to “fade.” Here also the intervals were found to be,

roughly speaking in arithmetical proportion. For similar reasons the

student is advised to review his “lecture-notes” shortly after taking.

them, and if possible, to review them again the evening of the same

day. Then the lapse of a week or two does not make nearly so much

difference. When once he has forgotten so much that the various asso-

ciations originally made have vanished, a considerable portion of the

material is irretrievably lost.

2. The Relation of the Length of Material to Time Taken for

Learning.—Tables were presented to show that the relation depended

almost wholly upon the division of the time spent in learning, 1. e., the

distribution of the time intervals. In other words, the relation, or ratio,

depends upon the method used in memorizing. Only three methods

were considered: The “continuous” or “mass” method; the once-per-day

method; and the once-per-week method. Up to a certain point, with

some individuals, when digits were used as material, the time varied

directly as the square of the number of digits, when the continuous

method was used. By the once-per-day method, however, the time varied,

roughly speaking, directly as the length of the material. It was shown

that in order to get the best results the same subject should take all

the various lengths of material used, and that it would be unfair to dis-

370 ANNALS NEW YORK ACADEMY OF SCIENCES

tribute the varying lengths among different subjects. As only one

_ method can be tried at a time, an experiment of this nature must needs

extend over a period of several years. In the case of prose, by the once-

per-day method, 500 words were memorized in as few days as the 95-

word passage. The time may therefore be said to vary directly as the

length of the passage. The same holds true for digits and nonsense-

syllables, but not to so great a degree; for the number of days needed

for 200 nonsense-syllables was considerably greater than that needed

for 20. By the “continuous” method, however, we observe that where

the 100-word passage was memorized in 9 minutes, the 500-word passage

took 52 minutes—nearly 6 times as much time being required, although

the passage is only 5 times as long. This is much more strikingly shown

when we examine the curve obtained for the digits. Here we see that

although it took only 5 minutes to learn 24 digits, it took 2 hours and

34 minutes to learn 200—more than 31 times as long instead of 8.

In short, it is obvious that the once-per-day method is—to say nothing

of giving a far superior retention—far more economical than the “con-

tinuous” method. This is especially so for material memorized by motor

associations such as nonsense-syllables or digits. }

The Section then adjourned.

F. Lyman WELLS,

Secretary.

BUSINESS MEETING.

6 May, 1912.

The Academy met at 8:15 p. mM. at the American Museum of Natural

History, President McMillin presiding.

The minutes of the last business meeting were read and approved.

The Recording Secretary reported the deaths of the following.

members : |

Col. John Jacob Astor, an Active Member for 18 years, lost with

the Titanic.

Mr. Isidor Straus, an Active Member for 6 years, lost with the

Titanic.

Col. John Weir, a Life Member for 5 years, lost with the Titanic.

Mr. George Borup, an Active Member for 4 months.

The Recording Secretary spoke of the great loss to the Academy,

the Museum and the scientific world at large entailed by the death of

Mr. Borup, who was to have been the leader of the Crocker Land Ex-

pedition organized under the auspices of the American Museum of

RECORDS OF MEETINGS Syl

Natural History and the American Geographical Society for the purpose

of arctic exploration during the years 1912-1914.

The Academy then adjourned.

EpmMuNpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

6 May, 1912.

Section met at 8:30 Pp. m., Vice-President Woodman presiding, about

150 members and visitors being present.

The following papers were read by title:

Neil E. Stevens, Nores on THE STRUCTURE AND GLACIATION OF OvER:

LOOK MouNrAIN.

George F. Kunz, THE GmM-BEARING PEGMATITES OF LOWER CALI-

FORNIA. ie

The meeting was then given over to the following public lecture:

Alfred H. Brooks, GrEoLogy anpD MINERAL ReEsouRCES OF ALASKA.

Mr. Brooks has been in charge of Alaskan exploration for the United

State Geological Survey for the last ten years or more. No one is more

intimately acquainted with the geography, geology and resources of that

country. The lecture was illustrated with lantern slides, maps and

charts.

The Section then adjourned.

CHARLES P. BERKEY,

Secretary.

LECTURE.

ry WW UNe ISLE

James F. B. Bowles: SANITATION OF THE PANAMA CANAL.

SECTION OF BIOLOGY.

13 May, 1912.

Section met at 8:15 Pp. m., Vice-President Lucas presiding.

The minutes of the last meeting of the Section were read and

approved. j

372 ANNALS NEW YORK ACADEMY OF SCIENCES

The following programme was then offered:

R. D. O. Johnson, Notr ON THE HABITS OF THE CLIMBING CAT-

FISH (Arges marmoratus) FROM THE UNITED

StTaTEs OF CoLomBi4A. (Read in abstract by

the Secretary.)

Bashford Dean, ON THE CHANGES IN THE BEHAVIOR OF THE

EEL (Conger malabaricus) DURING ITs TRANS-

FORMATION.

Bashford Dean, Do DrvELoPING EmBryos GivE REAL CLUES AS

To Lines oF DESCENT?

William K. Gregory, Norrs on CERTAIN PRINCIPLES OF QUADRUPEDAL

LocoMOTION AND ON THE MECHANISM OF THE

Limss oF Hoorep ANIMALS.

F. F. Hahn, On THE DictyonEMA Fauna oF Navy ISLAND,

3 New Brunswick. (Read by title.)

SUMMARY OF PAPERS.

Mr. Johnson’s paper is published as pages 327-333 of this volume.

Professor Dean said in abstract: When at Misaki, Japan, ihe speaker:

had made observations upon the structure and behavior of a living

leptocephalus larva which was kept alive in an aquarium for over three

weeks, during this time undergoing its metamorphosis. Especially in-

teresting is the rapidity with which the behavior of the young eel changes

from day to day in its methods of swimming and resting, response to

stimuli, ete. The speaker suggested that these marked differences in be-

havior in successive stages were correlated with kaleidoscopic changes in

elements of the central nervous system ; that when more fully known this

would probably afford a suggestive case of parallelism between psychic

reactions and neurological conditions. The paper was illustrated by

drawings and diagrams.

Professor Dean in his second paper said in abstract: After reviewing

the history of the question and touching upon the modern reaction

against the extreme views of Haeckel the speaker endeavored to show

that a comprehensive study of the anatomy and embryology of ganoid

and teleost fishes in the light of paleontological data gave strong evidence

in the affirmative.

The Secretary gave an abstract of a communication from Dr. P.

Bachmetjew, of Sofia, relating to the physiology of “Vesperugo pipi-

strellus’ and “Muiniopterus schreibersu.’ In some cases these bats

RECORDS OF MEETINGS 373

have been thawed out and the heart action had resumed even after the

body had been cooled to —7° Cent. below the body temperature.

The Section then adjourned.

Wititam K. GREGoRY,

Secretary.

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY.

20 May, 1912.

Section met at 8:15 P. m., in the Doremus Lecture Room of the

Chemistry Building of the College of the City of New York, Professor

Poor presiding.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered:

Charles Baskerville, TunGsTEN.

SUMMARY OF PAPER.

Professor Baskerville pointed out that tungsten at one time was

hardly mentioned in text books, but that now it is a substance of con-

siderable importance. It was discovered in tin-ore by Carl W. Scheele

in the year 1781. In 1848, tungsten salts were used for fixing colors in

cotton, and, in 1857, the fireproofing of draperies by tungsten was pro-

posed. Tungsten is used in making bronze and steel. Tungsten steel

retains its temper even when red hot and is better than the best carbon

steel known. The rims of car wheels are made of tungsten. steel.

The speaker then gave an interesting account of the invention and

development of tungsten lamps. He spoke at some length of the very

great practical difficulties that had to be overcome owing to the fact

that the tungsten filaments were brittle. Finally, however, this was

overcome so that now the tungsten incandescent lamp is the best one

on the market.

The Section then adjourned.

F. M. PEDERSEN,

Secretary.

BUSINESS MEETING.

% OcToBER, 1912.

The Academy met at 8:25 Pp. m. at the American Museum of Natural

History, President McMillin presiding.

374 ANNALS NEW YORK ACADEMY OF SCIENCES

The minutes of the last business meeting were read and approved.

The Recording Secretary reported the following deaths:

Ferdinand Zirkel, Honorary Member for 8 years.

Jules Henri Poincaré, Honorary Member for 12 years.

The Academy then adjonrned.

EpmMuND Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

7 OcToBER, 1912.

Section met at 8:30 p. M., Vice-President Woodman pres

35 members and visitors betta present.

The minutes of the last meeting of the Section were read and

approved.

Before the announced papers were called for the chairman asked

Professor J. F. Kemp to give some account of his recent trip to Panama.

A first-hand general account of his experiences and geologic observa-

tions was given. Questions were asked by members of the Section.

The following programme was then offered:

D. W. Johnson, THe Wesrwarp TRIP OF THE TRANSCONTINENTAL

EXCURSION OF THE AMERICAN GEOGRAPHICAL

SOcIETY. k

George H. Girty, Grotocic AGE oF THE BEDFORD SHALE OF OHIO.

SUMMARY OF PAPERS.

Dr. Johnson gave an interesting account of the make-up of the party,

the method of travel, the places of greater interest and some of the

special features to which most attention was given.

Dr. Girty’s paper was read in part by Professor Grabau and its bear-

ings were commented upon. It has been published as pages 295-319 of

this volume. |

The Section then adjourned.

CHARLES P. BERKEY,

Secretary.

RECORDS OF MEETINGS Bid)

SECTION OF BIOLOGY.

14 OctoBER, 1912.

~ Section met at 8:15 p. m., Vice-President Lucas presiding.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered:

Roy W. Miner, TypicaAL Marine INVERTEBRATE ASSOCIATION FROM

Woops Hotz To Casco Bay.

Roy C. Andrews, AN EXPLORATION OF NORTHEASTERN Korwa.

SUMMARY OF PAPERS.

Mr. Miner described and illustrated a series of typical invertebrate

faunal complexes or associations of the eastern Atlantic coast. He gave

views of many beautiful models and faunal groups which had been

mode for the American Museum, illustrating ecological relations and the

dominance of certain groups in particular localities.

Mr. Andrews gave an account of an exploration made for the Ameri-

can Museum of Natural History in a territory not hitherto studied by

zoologists.

The Section then adjourned.

WILLiAM K. GREGORY,

Becrey,

- SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY.

21 OcToBER, 1912.

Section.met at 8:30 Pp. M., Vice-President Poor presiding.

The minutes of the last meeting of the Section were read and

oparoret. .

Professor Charles Lane Poor was sonianted for Teen of

the Academy and Chairman of.the Section for 1913.

Professor F. M. Pedersen was elected Secretary.

The Committee on the future of the Section, consisting of Professors

Poor, Trowbridge and Pedersen then made its report, which was referred

to the Council for consideration.

F. M. PEDERSEN,

Secretary.

376 ANNALS NEW YORK ACADEMY OF SCIENCES

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

28 OcroBER, 1912.

Section met in conjunction with the American Ethnological Associa-

tion at 8:15 Pp. m., General James Grant Wilson presiding, about 128

members and visitors being present.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered :

Franz Boas, A YEAR IN MEXIco.

SUMMARY OF PAPER.

Professor Boas gave an outline of the work of archeological excavation

and linguistic research conducted by him during his stay in Mexico.

His lecture was illustrated by numerous stereopticon slides showing

especially pottery found in different layers in the Valley of Mexico.

The Section then adjourned.

F. LyMan WELLS,

Secretary. -

BUSINESS MEETING.

4 NOVEMBER, 1912.

The Academy met at 8:30 Pp. Mm. at the American Museum of Natural

History, Vice-President Woodman presiding.

The minutes of the last business meeting were read and approved.

The Recording Secretary reported the following death:

Morris Loeb, an Active Member and Fellow for 20 years.

The preparation of a suitable minute for the records of the Academy

was referred to Professor J. J. Stevenson and Dr. HE. O. Hovey.

Announcement was made from the Council of the engagement of

Dr. Alexis Carrel, to give an address before the Academy on 11 No-

vember, regarding his recent experimental work in physiology and of

Professor Hugo de Vries for an address upon experimental evolution

to be delivered 6 December.

The Academy then adjourned.

Epmunp Oris Hovey,

Recording Secretary.

RECORDS OF MEETINGS Birdy

SECTION OF GEOLOGY AND MINERALOGY.

+ NovEeMBER, 1912.

Section met al, 8:45 p. m., Vice-President Woodman presiding, 23

members and visitors being present.

In the absence of Secretary Berkey, Dr. Hovey was elected Secretary

pro tem.

The minutes of the last meeting of the Section were read and

approved.

On motion duly made and seconded, Professor J. E. Woodman was

nominated chairman of the Section and Vice-President of the Academy

for the year 1913.

Professor Berkey expressed the earnest wish that he be relieved of

the secretaryship which he had held five years. The Section acquiesced

in the request and passed a most cordial vote of thanks to him for his

long, faithful and efficient services, which have contributed so much to

the success of the Section. Professor C. T. Kirk of the Normal College

of the City of New York, was then nominated Secretary of the Section

for 1913 and unanimously elected.

The chairman then appointed Professor J. F. Kemp, Dr. George F.

Kunz and Dr. E. O. Hovey to serve with himself as a committee, in

response to a request of the Council, to consider the condition of the

Section and to make recommendations for its future work, this Com-

mittee to report to the Council.

The Secretary pro tem then read a letter from R. B. Earle of the

Department of Geology, New York University, asking for a grant of

$200 to assist him in carrying on research work on the origin and

history of certain types of interbedded iron ores. On motion, this

application was approved and referred to the Committee on Grants

from Research Funds for consideration.

The following programme was then offered:

F. S. Hintze, THE Fosstts AND HorIzON OF THE DRIFT

PEBBLES. :

Marjorie O’Connell, PRESENT OPINIONS ON THE HABITS OF TEE

EURYPTERIDS. .

A. W. Grabau, WAS THERE A FoRMER GoAT ISLAND AT NIAGARA

GLEN ?

Mr. Hintze’s paper was discussed by Professor A. W. Grabau, Pro-

fessor J. F. Kemp, Mr. B. E. Dodge and Professor D. 8. Martin.

Miss O’Connell’s paper was discussed by Professor Grabau.

378 ANNALS NEW YORK ACADEMY OF SCIENCES

SUMMARY OF PAPER.

Professor Grabau said in abstract: Foster’s Flat below the whirlpool

of Niagara and the topography and cross section of the gorge show —

that the falls of Niagara were once localized there in the same style

as is now the case at Goat Island. The same kind of development of

river work is well illustrated also at the falls of the Genesee. _

The paper was discussed by Professor D. W. Johnson, who cited what

is probably a similar case from near St. Anthony’s Falls, Minnesota,

of the Mississippi River.

The Section then adjourned.

: EpmMuND Otis Hovey,

Secretary pro tem.

SECTION OF BIOLOGY.

11 NovrempBer, 1912.

On this occasion the Section of Biology co-operated with the Academy

as a ‘whole and with the American Museum of Natural History in

welcoming Dr. Alexis Carrel, recipient of the Nobel Prize in medicine,

1912, who gave a lecture in the large auditorium, entitled “Results of

the Suture of Blood Vessels and the Transplantation of Organs,” about

800 persons being in attendance. After the lecture an informal re-

ception, attended by the officers, members and friends of the Academy

was held in honor of the lecturer.

. Witi1am K. Grecory,

Secretary.

SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY.

18 NovEMBER, 1912.

Section met at 8:15 Pp. M., Vice-President Poor presiding.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered:

Albert B. Pacini, THe DistrisuTion or FERRIC CHLORIDE BETWEEN

ETHER AND AQqurous Hyprocutoric AcID AT

25> ©.

Charles Lane Poor, THE Cause or THE TIDES.

RECORDS OF MEETINGS Bis)

SUMMARY OF PAPERS.

. Dr. Pacini gave a study of the distribution of ferric chloride under

the conditions which obtain in the use of Rothe’s method for the de-

termination of aluminum, nickel and other metals in steel, the mixed

chlorides in hydrochloric acid solution being shaken out with ether which

removes the greater portion of the ferric chloride. No decisive knowl-

edge was gained regarding the state of molecular aggregation of the

ferric chloride in the ether solution insomuch as data concerning the

degree of disassociation of ferric chloride in aqueous solution are not at

present available.

Graphical treatment of the constants obtained yields a curve of two

distinct sections: in low concentrations up to about 0.38 mols per liter

in the ether layer, a straight line; above this point a parabola satisfying

the equation (C, — .38)?— KC,, where K == -+ 1.8.

The application of the results to analytical separation lies in the

fact that the percentage of iron extracted from a hydrochloric acid

solution by shaking out with ether is greater relatively as the concen-

tration is lower, that is to say, the more dilute the original hydrochloric

acid solution of iron, the nearer complete is the extraction of ferric

chloride therefrom by ether.

Professor Poor gave a brief outline of the theories of the tides as

developed by La Place, Darwin and others, and contrasted these older

theories with recent investigations and theories of Dr. Harris, of the

United States Coast and Geodetic Survey. Until the researches of Dr.

Harris appeared, the tides were considered as a world phenomenon,

and primarily due to a large wave which originates in the Pacific Ocean

and travels around the world at varying speeds, due to the depth of the

oceans. This wave was supposed to take some fifty hours to travel from

the Pacific around Cape Horn to the shores of New York. Dr. Harris

considers the tides as purely local phenomena; the tide of each ocean

basin is primarily due to a standing wave or oscillation originating in

that basin and practically independent of the oscillations or tides in other

basins. The tides at New York and the Atlantic Coast, under this theory,

originate in the North Atlantic basin and are wholly independent of the

tides in the Pacific and Indian Oceans.

The Section then adjourned.

: F. M. PEDERSEN,

Secretary.

880 ANNALS NEW YORK ACADEMY OF SCIENCES

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY.

25 NOVEMBER, 1912.

Section met in conjunction with the New York Branch of the Ameri-

can Psychological Association, Dr. R. S. Woodworth acting as chairman.

The afternoon session was held at the Psychological Laboratory, Co-

lumbia University, at 4:10 Pp. m. and the evening session was held at

the American Museum of Natural History at 8:15 P. M.

The following programme was offered:

F. Krueger, DIFFERENT-TONES AND CONSONANCE.

Raymond Dodge, T'H= ATrempr TO MrasuRE MENTAL WoRK AS A

PsycHo-DyNAMIC PROCESS.

Robert M. Yerkes, THE PsycHoLocy oF THE EARTHWORM.

The Section then adjourned.

F. Lyman WELLS,

Secretary.

BUSINESS MEETING.

2 DECEMBER, 1912.

The Academy met at 8:15 Pp. M. at the American Museum of Natural

History, President Emerson McMillin presiding.

The following candidates for membership in the Academy, recom-

mended by Council, were duly elected:

ACTIVE MEMBERSHIP.

D. W. Johnson, Columbia University.

AssocIaATe MEMBERSHIP.

Marjorie O’Connell, 616 West 182nd Street.

The Academy then adjourned.

EpmuNnpD Otis Hovey,

Recording Secretary.

SECTION OF GEOLOGY AND MINERALOGY.

2 DECEMBER, 1912. ~

Section met at 8:20 p. M., President McMillin presiding.

The meeting was devoted to the following public lecture:

RECORDS OF MEETINGS 381

Harry Fielding Reid, THe SrISMOGRAPH AND Wat rt THAcHEs.

Professor Reid described the characteristics of the seismograph, which

has now been sufficiently perfected to record strong earthquakes occur-

ring at the antipodes. The revelations of the instrument and problems

awaiting solution were discussed and what happens at the time of an

earthquake was explained. The lecture was illustrated with lantern

slides.

The Section then adjourned.

CHarLes P. BEerKEY,

Secretary.

LECTURE.

6 DECEMBER, 1912.

(In co-operation with the American Museum of Natural History.)

Hugo de Vries, EXPERIMENTAL EVOLUTION.

SECTION OF BIOLOGY.

9 DECEMBER, 1912.

Section met at 8:15 p. M., Professor Henry Fairfield Osborn

presiding.

The minutes of the last meeting of the Section were read and

approved.

The following programme was then offered:

C.-E. A. Winslow, A Musrum or Livine Bacteria.

A. J. Goldfarb, Tue INFLUENCE oF THE NERVOUS SYSTEM UPON

GRowTH.

A. J. Goldfarb, A New Mernop or Fusinc Eces oF THE SAME

SPECIES.

SUMMARY OF PAPERS.

Dr. Winslow said in abstract: The American Museum of Natural

History is the first museum of its kind to recognize that the relation

between man and his microbic foes is fundamentally a problem in

natural history and a problem of such interest and importance as to

warrant the creation of a special Museum Department of Health. The

prime function of this department is of course to present in the form

of effective exhibits the main facts about the parasites which cause

382 ANNALS NEW YORK ACADEMY OF SCIENCES

disease, their life history, the conditions which favor their spread to

man, their relations to intermediate insect hosts and the means by which

mankind may be protected from their attacks. In connection with

this work of public exhibition there seemed to be a unique oppprtunity

for maintaining, as a sort of study collection, a museum of living

bacteria for the benefit of .working laboratories all over the country.

American bacteriologists have heretofore been compelled to send. to

Vienna for authentic stock cultures, and many important type strains

have been lost because the laboratories in which they were isolated had

no facilities for keeping them permanently under cultivation. The

authorities of the Museum were quick to appreciate the importance of

the public service that could thus be rendered to those engaged in

bacteriological teaching and research and early in 1911 endorsed the

establishment of such a collection and bureau for the distribution of

bacterial culture. A circular was sent out, to which the various labora-

tories quickly responded by sending in the cultures in their possession.

On December 1, 1912, the collection included 578 strains representing

374 different named types and including most of the important patho-

genic and non-pathogenic forms which have been definitely described.

During the period of somewhat less than two years, from January 1,

1911, to December 1, 1912, the laboratory distributed to 122 different

colleges and research laboratories of the United States and Canada

1700 different cultures, in every case without charge. It is the policy

of the Department to send cultures free to all teaching laboratories of

college and university grade and to all research laboratories, whether

cultures are sent to it in return or not. Many cultures have been called

for by teaching laboratories for use in their class work. The most im-

portant service the laboratory has been able to render, however, has

been in furnishing authentic cultures to investigators who have been

making a study of certain special groups and the published papers

which have resulted, in which various detailed characters of the museum.

types are described, of course greatly increase the value of the

collection.

The paper was discussed by Professor Osborn.

Dr. Goldfarb described a series of experiments upon certain annelid

worms (Amphinoma Lumbricus) which showed that the presence of the

central nervous system was not essential for growth and regeneration.

Dr. Goldfarb, in his second paper, described a method of fusing

embryos and larvae of the sea urchin (Toxopneustes). After fertilizing

the eggs they were placed in sea water to which enough 5 molecular

Na Cl was added to make a solution of 35 to 75 per cent. The eggs:

RECORDS OF MEETINGS 383

were left in this solution about six hours and were then transferred to

normal sea water. The unfused larvae floated to the surface, the

fused ones, which were obtained in great numbers, remained at or near

the bottom. :

The Section then adjourned.

WiLuiam K. GREGORY,

Secretary.

ANNUAL MEETING.

15 DEcEMBER, 1912.

_ The Academy met in Annual -Meeting on Monday, 16 December,

1912, at the Hotel Endicott, at the close of the annual dinner, President

McMillin presiding.

The minutes of the last Annual Meeting, 18 December, tei were

read and approved.

Reports were presented by the Corresponding Secretary, the Record-

ing Secretary, the Librarian and the Editor, all of which, on motion,

were ordered received and placed on file. They are published herewith.

The Treasurer’s report showed a net cash balance of $1,555.51 on

hand at the close of business, 30 November, 1912. On motion, this

report was received and referred to the Finance Committee for

auditing.

The following candidates for honorary membership and fellowship,

recommended by Council, were duly elected:

Honorary MEMBERS.

Jliya Metchnikof, Biologist and Bacteriologist, Pasteur Institute, Paris, _

France, presented by Dr. C.-E. A. Winslow.

Sir John Murray, Oceanographer, Edinburgh, Scotland, presented -

by Dr. F. A. Lucas.

Sho Watasé, Zodlogist, Imperial University of Tokio, Japan, pre-

sented by Prof. Henry E. Crampton.

Frank D. Adams, Geologist, McGill pe aeuNe Montreal, presented

by Prof. James F. Kemp.

George E. Hale, Astronomer, Solar Observatory, California, pre-

sented by Dr. John Tatlock. |

FELLOWS.

' Felix Arnold, M. D., 824 St. Nicholas Avenue, New York.

Dr. R. B. Earle, New York University, New York.

384. ANNALS NEW YORK ACADEMY OF SCIENCES

Prof. Jesse E. Hyde, School of Mining, Kingston, Ontario.

Prof. D. W. Johnson, Columbia University, New York.

Dr. A. B. Pacini, 275 West 140th Street, New York.

Mr. V. Stefansson, American Museum of Natural History, New York,

Dr. F. Lyman Wells, McLean Hospital, Waverley, Massachusetts.

The Academy then proceeded to the election of officers for the year

1912. The ballots prepared by the Council in accordance with the By-

laws were distributed. On motion it was unanimously voted that Dr.

Stevenson cast one ballot for the entire list nominated by the Council.

This was done and they were declared elected, more than the requisite

number of members and fellows entitled to vote being present:

President, EMERSON MoMILLIN. -

Vice-Presidents, J. EpmMunD WoopMaANn (Section of Geology and

Mineralogy), W. D. MarrHrw (Section of Biology), CHarLes LANE

Poor (Section of Astronomy, Physics and Chemistry), W. P. MontTacuE

(Section of Anthropology and Psychology).

Corresponding Secretary, Henry E. Crampron.

Recording Secretary, EpMunD Otis Hovey.

Treasurer, Henry L. DoHERTY.

Librarian, RatpH W. Tower.

Editor, EpMuND OT1Is Hovey. ~

Councilors (to serve 3 years), FrepERIc A. Lucas, R. S. WoopwortH.

Finance Committee, Herson McMiuin, Freprric 8S. Les, G. F.

KUNZ.

At the close of the elections, Mr. Emerson McMillin gave his address

as retiring President, in which, after reviewing the present condition

of the Academy as derived from conference with a large number of the

men who have long been active in carrying on its various lines of work,

he made several recommendations regarding plans which might be

adopted for enlarging the usefulness and interest of the organization and

its meetings.

Mr. V. Stefansson then gave a most interesting summary account

of the expedition which he and Dr. R. M. Anderson made along the

Arctic coast of western North America, from Point Barrow to Corona-

tion Gulf during the years 1908-1912 inclusiye. At the close of his

lecture, Mr. Stefansson outlined the plans of the second expedition which

he is now organizing for geographical and ethnological work on Victoria,

Banks and Prince Patricks Islands in the years 1913-1916 inclusive, and

indicated the manner in which his expedition and the Crocker Land

Expedition will supplement each other’s work.

The Academy then adjourned. Epmunp Otis Hovey,

Recording Secretary. .

RECORDS OF MEETINGS 385

REPORT OF THE CORRESPONDING SECRETARY.

We have lost by death during the past year the following Honorary

Members: |

Sir George H. Darwin, elected 1899.

Sir Joseph D. Hooker, elected 1907.

Franz Leydig, elected 1900.

M. Jules H. Poincaré, elected 1900.

Eduard Strasburger, elected 1908.

Prof. Ferdinand Zirkel, elected 1904.

and the following Corresponding Members:

Paul Schweitzer, elected 1867.

George Jarvis Brush, elected 1876.

There are at present upon our rolls 44 Honorary Members and 125

Corresponding Members.

Respectfully submitted,

Henry EH. CRAMPTON,

Corresponding Secretary.

REPORT OF THE RECORDING SECRETARY.

During the year 1912, the Academy held 8 business meetings and 26

sectional meetings, at which 65 ‘stated papers were presented as follows:

Section of Geology and Mineralogy, 24 papers; Section of Biology, 16

papers ; Section of Astronomy, Physics and Chemistry, 6 papers; Section

of Anthropology and Psychology, 19 papers.

_ Seven public lectures have been given at the American Museum to

the members of the Academy and the Affiliated Societies and their

friends, as follows:

“The Planet Mars.” By Edward E. Barnard. °

“Astronomy, Education and Culture.” By F. S. Archenhold.

“Geology and Mineral Resources of Alaska.” By Alfred H. Brooks.

“Sanitation of the Panama Canal.” By James F. B. Bowles.

“Results of the Suture of Blood Vessels and the Transplantation of

Organs.” By Alexis Carrel.

“The Seismograph and What it Teaches.” By Harry Fielding Reid.

“Experimental Evolution.” By Hugo de Vries.

At the present time, the membership of the Academy is 488, which

includes 468 Active Members, 22 of whom are Associate Members, 86

386 ANNALS NEW YORK ACADEMY OF SCIENCES

Fellows, 90 Life Members and 11 Patrons and 20 Non-resident Mem-

bers. There have been 10 deaths during the year, 12 resignations have

become effective, 7 names have been dropped from the roll for non-

payment of dues, 2 names have been transferred to the list of Non-

Resident Members. Twelve new members have been elected during

the year. As the membership of the Academy a year ago was 502,

there has been a net loss of 14 during the year 1912. Announcement is

made with regret of the loss by death of the following members:

Col. John Jacob Astor, Active Member for 17 years.

George Borup, Active Member for 4 months.

Charles F. Cox, Active Member for 36 years.

Prof. Morris Loeb, Active Member for 20 years.

William Pennington, Active Member for 6 years.

Edward Russ, Active Member for 5 years.

Prof. John B. Smith, Active Member for 5 years.

Isidor Straus, Active Member for 7 years.

James Terry, Life Member for 30 years.

Col. John Weir, Life Member for 6 years.

Respectfully submitted, —

EpMuND Otis Hovey,

Recording Secretary.

REPORT OF THE LIBRARIAN.

The accessions to the library of the New York Academy of Sciences

during the current year have been, by exchange and donation, 313

volumes and 1,670 numbers. Successful efforts have again been made

to complete imperfect files of publications and special acknowledg-

ments from the Academy are herewith extended to the Institutul Me-

teorologic al Romaniei for the presentation of nineteen volumes (1892—

1911) ; to the Verein fiir Erdkunde zu Leipzig for the presentation of

five volumes (1865-1871); to the Belfast Natural History and Philo-

sophical Society for the presentation of seven volumes (1872-1881) and

to the Société Nationale des Sciences Naturelles et Mathématiques de

Cherbourg for the presentation of six volumes (1858-1864). I am also

pleased to report that the books of the Academy have been more ex-

tensively used than during any of the preceding years.

Respectfully submitted,

RaLtpu W. TowEr, —

Labrarian.

RECORDS OF MEETINGS 397

REPORT OF THE EDITOR.

The Editor reports that during the past fiscal year there were issued

pages 177-263 completing Volume XXI and pages 1-337 of Volume

XXII.

Respectfully submitted, Epmunp Oris Hovey,

Editor.

REPORT OF THE TREASURER.

RECHIPTS.

DECEMBER 1, 1911—NovEMBER 30, 1912.

Cashvon hand December 1 19Id a, Se ooh. ce Soe chew eeew cae $1,356.58

MEME MPeLShip: KCCSsc eres sis sre ciao rere er eee lade axkiqee ors we eels 200.00

Income from investments:

Interest on mortgages on New York City real estate.. $860.00

Interest on railroad and other bonds................ 1,375 .00

2,235.00

MMKELES FeO mes aia BATA CAG lars snsvene aise el oa iets See dns ee: 4% (eile coile) io -elrews whew euaneuseaue 48.29

Active membership dues, 1909...............20.. cee eeee $10.00

ff on Ce yA ret ee nn Opie ncarec Re nOn Gig Ss CRO ORE Roe 50.00

t me HO SION Lees Gagan ck ecceeci orctoeieaa isc ccna 245 .00

:; seamen T VINA anes crar terra haya! Sia S ences we feve tat ois 3,185.00

3,490.00

Associate membership dues, 1911..................2005- $6.00

se s Sra aan Dons nrekiay a iusnaveye creche SeMa esc 45.00

51.00

SALE SMO ses CALTOMS rapid ten seedoyevel Sik cieisiccei visieiene aise aie-ateersloieecmens 89.30

SUUSCLIPLONS MOPAMMUAlGINMeM see cases oe ce cles cle meee 178.00

PIU ete peter ewag eect ieiencuater te rehcirolel cers iesei sis cir: eleeiene/siae sve siotaileualacevs)s $7,648.17

DISBURSEMENTS.

DECEMBER 1, 1911—-NoVEMBER 30, 1912.

Publications on account of Annals... .. 5... 06cc.. ss ccenes corre see $1,511.01

IP MONCTON OIE. JVM Mee BOG B60 Boe ee Cae ce eons 597.75

Recording Secretary’S CXPeEMSES........ 0.00 c cee cece ee eee eee eeee 298 .55

Recording Secretary’s and Hditor’s allowance...........-.20.+-s. 1,200.00

Lecture Committee........ SAIN horse Nayar ateai'd ail, cig ates cleners Sue ateseens 200.00

GE CME AGSX CH SOSH eee a Hater reese neilalievorcine sais evenareitrsiw is nceseie sie veihs eselions 300.65

Esther Herrman Research Fund corte) AR ya aiage eteteleae Pasahs lata: sieneat ete c ete 800.00

Interest charge on bond purchased.............--2.eeeeceecceeeee 5.00

Section of Geology and Mineralogy.............2eeeeeeeeeeceeeees 25.00

Cals Ro ripe lie leprae ar iey oe, LisaNova er sis Sidley etsie 6) Silsie oiaiotaris elev e alesis) saier evets 1,555.51

888 ANNALS NEW YORK ACADEMY OF SCIENCES

BALANCE SHEET, NOVEMBER 30, 1912.

Investments (cost)........ $42,631.25 Permanent Fund.......... $22,812.57

Cash on hand............. 1,555.51 Publication Fund.......... 3,000.00

Audubon Fund............ 2,500.00

Hsther Herrman Research

BUG ase hess hele ta ee 10,000.00

John Strong Newberry

Ah) 010 eee IER IGS 6 0.6 1,000.00

Income Permanent Fund... 2,624.32

Income Audubon Fund..... 334.75

Income Hsther Herrman

D000 Wat erae rere Hi als iS craic 1,465.96

Income Newberry Fund.... 449.16

$44,186.76 $44,186.76

Henry L. DOHERTY,

Treasurer.

8 JANUARY, 1913.

Examined and found to be correct,

GEORGE F. KUNZ,

FREDERIC S. LEE,

Auditing Committee.

THE ORGANIZATION OF THE NEW YORK ACADEMY OF

SCIENCES

THE ORIGINAL CHARTER

AN ACT TO INCORPORATE THE

LYCEUM OF NATURAL HISTORY IN THE CITY OF NEW YORK

Passed A pril 20, 1818

WHerzEAS, The members of the Lyceum of Natural History have peti-

tioned for an act of incorporation, and the Legislature, impressed with the

importance of the study of Natural History, as connected with the wants,

the comforts and the happiness of mankind, and conceiving it their duty

to encourage all laudable attempts to promote the progress of science in

this State—therefore, é:

1. Be tt enacted by the People of the State of New York represented in

Senate and Assembly, That Samuel L. Mitchill, Casper W. Eddy, Fred-

erick C. Schaeffer, Nathaniel Paulding, William Cooper, Benjamin P.

Kissam, John Torrey, William Cumberland, D’Jurco V. Knevels, James

Clements and James Pierce, and such other persons as now are, and may

from time to time become members, shall be, and hereby are constituted a

body corporate and politic, by the name of Lyceum oF Natura History

IN THE City oF New York, and that by that name they shall have per-

petual succession, and shall be persons capable of suing and being sued,

pleaded and being impleaded, answering and being answered unto, de-

fending and being defended, in all courts and places whatsoever ; and may

have a common seal, with power to alter the same from time to time; and

shall be capable of purchasing, taking, holding, and enjoying to them and

their successors, any real estate in fee simple or otherwise, and any goods,

chattels, and personal estate, and of selling, leasing, or otherwise dispos-

ing of said real or personal estate, or any part thereof, at their will and

pleasure: Provided always, that the clear annual value or income of such

real or personal estate shall not exceed the sum of five thousand dollars:

Provided, however, that the funds of the said Corporation shall be used

and appropriated to the promotion of the objects stated in the preamble

to this act, and those only.

2. And be wt further enacted, That the said Society shall from time to

time, forever hereafter, have power to make, constitute, ordain, and estab-

lish such by-laws and regulations as they shall judge proper, for the elec-

(389)

390 ANNALS NEW YORK ACADEMY OF SCIENCES

tion of their officers; for prescribing their respective functions, and the

mode of discharging the same; for the admission of new members; for the —

government of the officers and members thereof; for collecting annual

contributions from the members towards the funds thereof; for regulat-

ing the times and places of meeting of the said Society; for suspending

or expelling such members as shall neglect or refuse to comply with the

by-laws or regulations, and for the managing or directing the affairs and

concerns of the said Society: Provided such by-laws and regulations be

not repugnant to the Constitution and laws of this State or of the United

States.

3. And be tt further enacted, That the officers of the said Society shall

consist of a President and two Vice-Presidents, a Corresponding Secre-

tary, a Recording Secretary, a Treasurer, and five Curators, and such

other officers as the Society may judge necessary; who shall be annually

chosen, and who shall continue in office for one year, or until others be

elected in their stead; that if the annual election shall not be held at any

of the days for that purpose appointed, it shall be lawful to make such

election at any other day; and that five members of the said Society,

assembling at the place and time designated for that purpose by any by-

law or regulation of the Society, shall constitute a legal meeting thereof.

4. And be it further enacted, That Samuel L. Mitchill shall be the

President; Casper W. Eddy the First Vice-President; Frederick C.

Schaeffer the Second Vice-President; Nathaniel Paulding, Correspond-

ing Secretary; William Cooper, Recording Secretary; Benjamin P. Kis-

sam, Treasurer, and John Torrey, William Cumberland, D’Jurco V.

Knevels, James Clements, and James Pierce, Curators; severally to be

the first officers of the said Corporation, who shall hold their respective

offices until the twenty-third day of February next, and until others shall

be chosen in their places.

5. And be it further enacted, That the present Constitution of the said

Association shall, after passing of this Act, continue to be the Constitu-

tion thereof; and that no alteration shall be made therein, unless by a

vote to that effect of three-fourths of the resident members, and upon the

request in writing of one-third of such resident members, and submitted

at least one month before any vote shall be taken thereupon.

State of New York, Secretary's Office.

I certiry the preceding to be a true copy of an original Act of the

Legislature of this State, on file in this Office.

ARCH’D CAMPBELL,

ALBANy, April 29, 1818. Dep. Sec’y.

ORGANIZATION | 39]

ORDER OF COURT

ORDER OF THE SUPREME COURT OF THE STATE OF NEW YORK

TO CHANGE THE NAME OF

THE LYCEUM OF NATURAL HISTORY IN THE CITY OF

NEW YORK

THE NEW YORK ACADEMY OF SCIENCES

WHEREAS, in pursuance of the vote and proceedings of this Corpora-

tion to change the corporate name thereof from “The Lyceum of Natural

History in the City of New York” to “The New York Academy of Sci-

ences,” which vote and proceedings appear to record, an application has

been made in behalf of said Corporation to the Supreme Court of the

State of New York to legalize and authorize such change, according to

the statute in such case provided, by Chittenden & Hubbard, acting as

. the attorneys of the Corporation, and the said Supreme Court, on the 5th

day of January, 1876, made the following order upon such application in

the premises, viz:

At a special term of the Supreme

Court of the State of New York,

held at the Chambers thereof, in

the County Court House, in the

City of New York, the 5th day

of January, 1876:

Present—Hon. Gro. C. BARRETT, Justice.

In the matter of the application of

the Lyceum of Natural History

in the City of New York to au-

thorize it to assume the corporate

name of the New York Academy

of Sciences.

On reading and filing the petition of the Lyceum of Natural History

in the City of New York, duly verified by John S. Newberry, the Presi-

dent and chief officer of said Corporation, to authorize it to assume the

corporate name of the New York Academy of Sciences, duly setting forth

392 ANNALS NEW YORK ACADEMY OF SCIENCES

the grounds of said application, and on reading and filing the affidavit of

Geo. W. Quackenbush, showing that notice of such application had been

duly published for six weeks in the State paper, to wit, The Albany

Evening Journal, and the affidavit of David 8S. Owen, showing that notice

of such application has also been duly published in the proper newspaper

of the County of New York, in which county said Corporation had its

business office, to wit, in The Daily Register, by which it appears to my

satisfaction that such notice has been so published, and on reading and

filing the affidavits of Robert H. Browne and J. 8. Newberry, thereunto

annexed, by which it appears to my satisfaction that the application is

made in pursuance of a resolution of the managers of said Corporation to

that end named, and there appearing to me to be no reasonable objection

to said Corporation so changing its name as prayed in said petition: Now

on motion of Grosvenor 8. Hubbard, of Counsel for Petitioner, it is

Ordered, That the Lyceum of Natural History in the City of New

York be and is hereby authorized to assume the corporate name of The

New York Academy of Sciences. :

Indorsed: Filed January 5, 1876,

: A copy. Wm. WatsH, Clerk.

Resolution of THE ACADEMY, accepting the order of the Court, passed

February 21, 1876

And whereas, The order hath been published as therein required, and

all the proceedings necessary to carry out the same have been had, There-

fore:

Resolved, That the foregoing order be and the same is hereby accepted

and adopted by this Corporation, and that in conformity therewith the

corporate name thereof, from and after the adoption of the vote and reso-

lution herein above referred to, be and the same is hereby declared to be

THE NEW YORK ACADEMY OF SCIENCES.

AMENDED CHARTER

Marcy 19, 1902

CHAPTER 181 oF THE Laws oF 1902

Aw Act to amend chapter one hundred and ninety-seven of the laws of

eighteen hundred and eighteen, entitled “An act to incorporate the Ly-

ceum of Natural History in the City of New York,’ a Corporation now

known as The New York Academy of Sciences and to extend the powers

of said Corporation. .

ORGANIZATION 393

(Became a law March 19, 1902, with the approval of the Governor.

Passed, three-fifths being present.)

The People of the State of New York, represented in Senate and As-

sembly, do enact as follows:

Section I. The Corporation incorporated by chapter one hundred

and ninety-seven of the laws of eighteen hundred and eighteen, entitled

“An act to incorporate the Lyceum of Natural History in the City of

New York,” and formerly known by that name, but now known as The

New York Academy of Sciences through change of name pursuant to

order made by the supreme court at the city and county of New York, on

January fifth, eighteen hundred and seventy-six, is hereby authorized and

_ empowered to raise money for, and to erect and maintain, a building in

the city of New York for its use, and in which also at its option other

scientific societies may be admitted and have their headquarters upon

such terms as said Corporation may make with them, portions of which

building may be also rented out by said Corporation for any lawful uses

for the purposes of obtaining income for the maintenance of such build-

ing and for the promotion of the objects of the Corporation ; to establish,

own, equip, and administer a public library, and a museum having es-

pecial reference to scientific subjects; to publish communications, trans-

actions, scientific works, and periodicals; to give scientific instruction by

lectures or otherwise; to encourage the advancement of scientific research

and discovery, by gifts of money, prizes, or other assistance thereto. The

building, or rooms, of said Corporation in the City of New York used

exclusively for library or scientific purposes shall be subject to the pro-

visions and be entitled to the benefits of subdivision seven of section four

of chapter nine hundred and eight of the laws of eighteen hundred and

ninety-six, as amended.

Section II. The said Corporation shall from time to time forever

hereafter have power to make, constitute, ordain, and establish such by-

laws and regulations as it shall judge proper for the election of its officers ;

for prescribing their respective functions, and the mode of discharging

the same; for the admission of new members; for the government of offi-

cers and members thereof; for collecting dues and contributions towards

the funds thereof; for regulating the times and places of meeting of said

Corporation ; for suspending or expelling such members as shall neglect

or refuse to comply with the by-laws or regulations, and for managing or.

directing the affairs or concerns of the said Corporation: and may from

time to time alter or modify its constitution, by-laws, rules, and regula-

tions.

394. ANNALS NEW YORK ACADEMY OF SCIENCES

Section III. The officers of the said Corporation shall consist of a

president and two or more vice-presidents, a corresponding secretary, a

recording secretary, a treasurer, and such other officers as the Corporation

may judge necessary; who shall be chosen in the manner and for the

terms prescribed by the constitution of the said Corporation.

Srction IV. The present constitution of the said Corporation shall,

after the passage of this act, continue to be the constitution thereof until

amended as herein provided. Such constitution as may be adopted by a

vote of not less than three-quarters of such resident members and fellows

of the said New York Academy of Sciences as shall be present at a meet-

ing thereof, called by the Recording Secretary for that purpose, within

forty days after the passage of this act, by written notice duly mailed,

postage prepaid, and addressed to each fellow and resident member at

least ten days before such meeting, at his last known place of residence,

with street and number when known, which meeting shall be held within

three months after the passage of this act, shall be thereafter the consti-

tution of the said New York Academy of Sciences, subject to alteration

or amendment in the manner provided by such constitution.

Suction V. The said Corporation shall have power to consolidate, to

unite, to co-operate, or to ally itself with any other society or association

in the city of New York organized for the promotion of the knowledge or

the study of any science, or of research therein, and for this purpose to

receive, hold, and administer real and personal property for the uses of

such consolidation, union, co-operation, or alliance subject to such terms

and regulations as may be agreed upon with such associations or societies.

Section VI. This act shall take effect immediately.

STATE OF New YORK,

OFFICE OF THE SECRETARY OF STATE.

I have compared the preceding with the original law on file in this

office, and do hereby certify that the same is a correct transcript there-

from, and the whole of said original law.

Given under my hand and the seal of office of the Secretary of State,

at the city of Albany, this eighth day of April, in the year one thousand

nine hundred and two.

JoHN T. McDonoucH, |

Secretary of State.

ORGANIZATION 395

CONSTITUTION

-ApopTEeD, APRIL 24, 1902, AND AMENDED AT SUBSEQUENT TIMES

ArticLE I. The name of this Corporation shall be The New York

Academy of Sciences. Its object shall be the advancement and diffusion

of scientific knowledge, and the center of its activities shall be in the City

of New York.

ArticLE II. The Academy shall consist of five classes of members,

namely: Active Members, Fellows, Associate Members, Corresponding

Members and Honorary Members. Active Members shall be the members

of the Corporation who live in or near the City of New York, or who,

having removed to a distance, desire to retain their connection with the

Academy. Fellows shall be chosen from the Active Members in virtue of

their scientific attainments. Corresponding and Honorary Members shall

be chosen from among persons who have attained distinction in some

branch of science. The number of Corresponding Members shall not

exceed two hundred, and the number of Honorary Members shall not

exceed fifty.

-ARTICLE III. None but Fellows and Active Members who have paid

their dues up to and including the last fiscal year shall be entitled to vote

or to hold office in the Academy.

' ArticLtE IV. The officers of the Academy shall be a President, as

many Vice-Presidents as there are sections of the Academy, a Correspond-

ing Secretary, a Recording Secretary, a Treasurer, a Librarian, an Editor,

six elected Councilors and one additional Councilor from each allied

society or association. ‘The annual election shall be held on the third

Monday in December, the officers then chosen to take office at the first

meeting in January following.

There shall also be elected at the same time a Finance Committee of

three.

ARTICLE V. ‘The officers named in Article IV shall constitute a Coun-

cil, which shall be the executive body of the Academy with general control

over its affairs, including the power to fill ad interim any vacancies that

may occur in the offices. Past Presidents of the Academy shall be ez-

officio members of the Council.

ARTICLE VI. Societies organized for the study of any branch of

science may become allied with the New York Academy of Sciences by

consent of the Council. Members of allied societies may become Active

Members of the Academy by paying the Academy’s annual fee, but as

396 ANNALS NEW YORK ACADEMY OF SCIENCES

members of an allied society they shall be Associate Members with the

rights and privileges of other Associate Members, except the receipt of

its publications. Each allied society shall have the right to delegate one

of its members, who is also an Active Member of the Academy, to the

Council of the Academy, and such delegate shall have all the rights and

privileges of other Councilors.

ARTICLE VII. The President and Vice-Presidents shall not be eligible

to more than one re-election until three years after retiring from office;

the Secretaries and Treasurer shall be eligible to re-election without

limitation. The President, Vice-Presidents and Secretaries shall be Fel-

lows. The terms of office of elected Councilors shall be three years, and

these officers shall be so grouped that two, at least one of whom shall be a

Fellow, shall be elected and two retired each year. Councilors shall not

be eligible to re-election until after the expiration of one year.

ArTICLE VIII. The election of officers shall be by ballot, and the can-

didates having the greatest number of votes shall be declared duly elected.

ARTICLE [X. Ten members, the majority of whom shall be Fellows,

shall form a quorum at any meeting of the Academy at which business is

transacted.

ARTICLE X. The Academy shall establish by-laws, and may amend

them from time to time as therein provided.

ARTICLE XI. This Constitution may be amended by a vote of not less

than three-fourths of the fellows and three-fourths of the active members

present and voting at a regular business meeting of the Academy, pro-

vided that such amendment shall be publicly submitted in writing at the

preceding business meeting, and provided also that the Recording Secre-—

tary shall send a notice of the proposed amendment at least ten days

before the meeting, at which a vote shall be taken, to each Fellow and

Active Member entitled to vote.

BY-LAWS

As ADOPTED, OCTOBER 6, 1902, AND AMENDED AT SUBSEQUENT TIMES

CHapTer I

OFFICERS

1. President. It shall be the duty of the President to preside at the

business and special meetings of the Academy; he shall exercise the cus-

tomary duties of a presiding officer.

2. Vice-Presidents. In the absence of the President, the senior Vice-

President, in order of Fellowship, shall act as the presiding officer.

sae

ORGANIZATION 397

3. Corresponding Secretary. The Corresponding Secretary shall keep

a corrected list of the Honorary and Corresponding Members, their titles

and addresses, and shall conduct all correspondence with them. He shall

make a report at the Annual Meeting.

4. Recording Secretary. The Recording Secretary shall keep the

minutes of the Academy proceedings; he shall have charge of all docu-

ments belonging to the Academy, and of its corporate seal, which he shall

affix and attest as directed by the Council; he shall keep a corrected list

of the Active Members and Fellows, and shall send them announcements

of the Meetings of the Academy ; he shall notify all Members and Fellows

of their election, and committees of their appointment; he shall give

notice to the Treasurer and to the Council of matters requiring their

action, and shall bring before the Academy business presented by the

Council. He shall make a report at the Annual Meeting.

5. Treasurer. The Treasurer shall have charge, under the direction

of the Council, of all moneys belonging to the Academy, and of their

investment. He shall receive all fees, dues and contributions to the

Academy, and any income that may accrue from property or investment ;

he shall report to the Council at its last meeting before the Annual Meet-

ing the names of members in arrears; he shall keep the property of the

Academy insured, and shall pay all debts against the Academy the dis-

charge of which shall be ordered by the Council. He shall report to the

Council from time to time the state of the finances, and at the Annual

Meeting shall report to the Academy the receipts and expenditures for

the entire year.

6. Librarian. The Librarian shall have charge of the library, under

the general direction of the Library Committee of the Council, and shall

conduct all correspondence respecting exchanges of the Academy. He

‘shall make a report on the condition of the library at the Annual Meeting. |

%. Editor. The editor shall have charge of the publications of the

Academy, under the general direction of the Publication Committee of

the Council. He shall make a report on the condition of the publications

at the Annual Meeting.

CHAPTER II

COUNCIL

1. Meetings. The Council shall meet once a month, or at the call of

the President. It shall have general charge of the affairs of the Academy.

2. Quorum. Five members of the Council shall constitute a quorum.

3. Officers. The President, Vice-Presidents and Recording Secretary

of the Academy shall hold the same offices in the Council.

398 ANNALS NEW YORK ACADEMY OF SCIENCES

4. Committees. The Standing Committees of the Council shall be:

(1) an Executive Committee consisting of the President, Treasurer, and

Recording Secretary; (2) a Committee on Publication; (3) a Committee

on the Library, and such other committees as from time to time shall be

authorized by the Council. The action of these committees shall be sub-

ject to revision by the Council.

Cuapter III

FINANCE COMMITTEE

The Finance Committee of the Academy shall audit the Annual Report

of the Treasurer, and shall report on financial questions whenever called

upon to do so by the Council. |

CHaAptTerR IV

ELECTIONS

1. Active Members. (a) Active Members shall be nominated in writ-

ing to the Council by at least two Active Members or Fellows. If ap-

proved by the Council, they may be elected at the succeeding business

meeting.

(6) Any Active Member who, eine removed to a distance ise the

city of New York, shall nevertheless express a desire to retain his connec-

tion with the Academy, may be placed by vote of the Council on a list of

Non-Resident Members. Such members shall relinquish the full privi-

leges and obligations of Active Members. (Vide Chapters V and X.)

2. Associate Members. Workers in science may be elected to Associate

Membership for a period of two years in the manner prescribed for Active ~

Members. They shall not have the power to vote and shall not be eligible

to election as Fellows, but may receive the publications, At any time sub-

sequent to their election they may assume the full privileges of Active

Members by paying the dues of such Members.

3. Fellows, Corresponding Members and Honorary Members. Nomi-

nations for Fellows, Corresponding Members and Honorary Members

may be made in writing either to the Recording Secretary or to the

Council at its meeting prior to the Annual Meeting. If approved by the

Council, the nominees shall then be balloted for at the Annual Meeting.

4, Officers. Nominations for Officers, with the exception of Vice-

Presidents, may be sent in writing to the Recording Secretary, with the

name of the proposer, at any time not less than thirty days before the

Annual Meeting. Hach section of the Academy shall nominate a candi-

ORGANIZATION 399

date for Vice-President, who, on election, shall be Chairman of the sec-

tion ; the names of such nominees shall be sent to the Recording Secretary

properly certified by the sectional secretaries, not less than thirty days

before the Annual Meeting. The Council shall then prepare a list which

shall be the regular ticket. This list shall be mailed to each Active Mem-

ber and Fellow at least one week before the Annual Meeting. But any

Active Member or Fellow entitled to vote shall be entitled to prepare and

vote another ticket.

CHAPTER V

DUES

1. Dues. The annual dues of Active Members and Fellows shall be

$10, payable in advance at the time of the Annual Meeting; but new

members elected after May 1, shall pay $5 for the remainder of the fiscal

year. .

The annual dues of elected Associate Members shall be $3, payable in

advance at the time of the Annual Meeting.

Non-Resident Members shall be exempt from dues, so long as they shall

relinquish the privileges of Active Membership. (Vide Chapter X.)

2. Members in Arrears. If any Active Member or Fellow whose dues

remain unpaid for more than one year, shall neglect or refuse to pay the

same within three months after notification by the Treasurer, his name

may be erased from the rolls by vote of the Council. Upon payment of

his arrears, however, such person may be restored to Active Membership

or Fellowship by vote of the Council.

3. Renewal of Membership. Any Active Member or Fellow who shall

resign because of removal to a distance from the city of New York, or

any Non-Resident Member, may be restored by vote of the Council to

Active Membership or Fellowship at any time upon application.

CHAPTER VI

PATRONS, DONORS AND LIFE MEMBERS

1. Patrons. Any person contributing at one time $1,000 to the general

funds of the Academy shall be a Patron and, on election by the Council,

shall enjoy all the privileges of an Active Member.

2. Donors. Any person contributing $50 or more annually to the

general funds of the Academy shall be termed a Donor and, on election

by the Council, shall enjoy all the privileges of an Active Memter.

3. Life Members. Any Active Member or Fellow contributing at one

time $100 to the general funds of the Academy shall be a Life Member

400 ANNALS NEW YORK ACADEMY OF SCIENCES

and shall thereafter be exempt from annual dues; and any Active Mem-

ber or Fellow who has paid annual dues for twenty-five years or more

may, upon his written request, be made a life member and be exempt

from further payment of dues.

CHaptTer VII

SECTIONS

1. Sections. Sections devoted to special branches of Science may be

established or discontinued by the Academy on the recommendation of

the Council. The present sections of the Academy are the Section of

Astronomy, Physics and Chemistry, the Section of Biology, the Section

of Geology and Mineralogy and the Section of Anthropology and Psy-

chology.

2. Organization. Tach section of the Academy shall have a Chairman

and a Secretary, who shall have charge of the meetings of their Section.

The regular election of these officers shall take place at the October or

November meeting of the section, the officers then chosen to take office at

the first meeting in January following.

3. Affiliation. Members of scientific societies affiliated with the

Academy, and members of the Scientific Alliance, or men of science intro-

duced by members of the Academy, may attend the meetings and present

papers under the general regulations of the Academy.

CHapter VIII

MEETINGS

1. Business Meetings. Business meetings of the Academy shall be

held on the first Monday of each month from October to May inclusive.

2. Sectional Meetings. Sectional meetings shall be held on Monday

evenings from October to May inclusive, and at such other times as the

Council may determine. The sectional meeting shall follow the business

meeting when both occur on the same evening.

3. Annual Meeting. The Annual Meeting shall be held on the third

Monday in December.

4. Special Meetings. A special meeting may be called by the Council,

provided one week’s notice be sent to each Active Member and Fellow,

stating the object of such meeting.

ORGANIZATION 401

CHAPTER IX

ORDER OF BUSINESS

1. Business Meetings. The following shall be the order of procedure

at business meetings: .

1. Minutes of the previous business meeting.

2. Report of the Council.

3. Reports of Committees.

4, Elections.

5. Other business.

2. Sectional Meetings. The following shall be the order of procedure

at sectional meetings:

1. Minutes of the preceding meeting of the section.

2. Presentation and discussion of papers.

3. Other scientific business.

3. Annual Meetings. The following shall be the order of procedure

at Annual Meetings: |

1. Annual reports of the Corresponding Secretary, Recording

Secretary, Treasurer, Librarian, and Editor.

2. Hlection of Honorary Members, Corresponding Members, and

Fellows.

Election of officers for the ensuing year.

4. Annual address of the retiring President.

CHAPTER X

PUBLICATIONS

1. Publications. The established publications of the Academy shall

be the Annals and the Memoirs. They shall be issued by the Editor

under the supervision of the Committee on Publications.

2. Distribution. One copy of all publications shall be sent to each

Patron, Life Member, Active Member and Fellow; provided, that upon

inquiry by the Editor such Members or Fellows shall signify their desire

to receive them. ;

3. Publication Fund. Contributions may be received for the publica-

tion fund, and the income thereof shall be applied toward defraying the

expenses of the scientific publications of the Academy.

402 ANNALS NEW YORK ACADEMY OF SCIENCES

CHAPTER XT

GENERAL PROVISIONS

1. Debts. No debts shall be incurred on behalf of the Academy, unless

authorized by the Council.

2. Bills. All bills submitted to the Council must be certified as to

correctness by the officers incurring them.

3. Investments. All the permanent funds of the ‘Weadete shall be

invested in United States or in New York State securities or in first

mortgages on real estate, provided they shall not exceed sixty-five per

cent. of the value of the property, or in first-mortgage bonds of corpora-

tions which have paid dividends continuously on their common stock for

a period of not less than five years, All income from patron’s fees, life-

membership fees and donor’s fees shall be added to the permanent fund.

4. Expulsion, etc. Any Member or Fellow may be censured, sus-

pended or expelled, for violation of the Constitution or By-Laws, or for

any offence deemed sufficient, by a vote of three-fourths of the Members

and three-fourths of the Fellows present at any business meeting, provided

such action shall have been recommended by the Council at a previous

business meeting, and also, that one month’s notice of such recommenda-

tion and of the offence charged shall have been given the Member accused.

5. Changes in By-Laws. No alteration shall be made in these By-

Laws unless it shall have been submitted publicly in writing at a business

meeting, shall have been entered on the Minutes with the names of the

Members or Fellows proposing it, and shall be adopted by two-thirds of

the Members and Fellows present and voting at a subsequent business

meeting.

MEMBERSHIP OF THE

NEW YORK ACADEMY OF SCIENCES

HONORARY MEMBERS

31 DECEMBER, 1912.

ELECTED.

1912. Frank D. Apams, Montreal, Canada.

1898. ArtTHuR Auwenrs, Berlin, Germany.

1889. CHARLES Barros, Lille, France.

1907. Wrtu1am Bateson, Cambridge, England.

1910. THropor Boveri, Wirzburg, Germany.

1901. CHaRLES VERNON Boys, London, England.

1904. W. C. Broécer, Christiana, Norway.

1911. Herrmann Crepner, Leipzig, Germany.

1876. W. Boyp Dawkins, Manchester, England.

1902. Sir JAmes Dewar, Cambridge, England. °

1901: Emit Fiscuer, Berlin, Germany.

1876. Sir Arcuipatp Gurxre, Haslemere, Surrey, England.

1901. James Gerxiz, Edinburgh, Scotland.

1898. Sir Davip Grux, London, England.

1909. K. F. GéseLt, Munich, Germany.

1889. GrorcE LINCOLN GoopaLE, Cambridge, Mass.

1909. PauL von GrotH, Munich, Germany. ’

1894. Ernst HAcKEL, Jena, Germany.

1912. Gzorce EH. Hate, Mt. Wilson, Calif.

1899. JuLius Hann, Vienna, Austria.

1898. GrorcE W. Hitt, West Nyack, N. Y.

1896. Amsrosius A. W. Husrecut, Utrecht, Netherlands.

1896. Ferix Kiser, Gottingen, Germany.

1909. Atrrep Lacrorx, Paris, France.

1876. ViKToR von Lane, Vienna, Austria.

1898. EH. Ray Lanxester, London, England.

1880. Sir Norman Lockyer, London, England.

1911. Ernst Macu, Vienna, Austria.

1912. Intya Metrounikor, Paris, France,

1912. Sir Jonw Murray, Edinburgh, Scotland.

1898. Friptyor Nansen, Christiana, Norway.

1908. WILHELM OstTWALD, Gross-Bothen, Germany.

1898. ALBRECHT PxENcK, Berlin, Germany.

(403)

404 ANNALS NEW YORK ACADEMY OF SCIENCES

wee

ELECTED.

1898. WILHELM PFEFFER, Leipzig, Germany.

1900. EpwarpD CHARLES PICKERING, Cambridge, Mass.

1911. Epwarp BacnaLu Poutton, Oxford, England.

1901. Sir Witu1Am Ramsay, London, England.

1899. Lord RayLereH, Witham, Essex, England.

1898. Hans H. Reuscu, Christiania, Norway.

1887. Sir Henry Enrretp Roscoz, London, England.

1887. HzrtnricH Rosensuscu, Heidelberg, Germany.

1912. SHo Warass, Tokyo, Japan.

1904. KARL VON DEN STEINEN, Berlin, Germany.

1896. JosEPpH JoHN THomson, Cambridge, England.

1900. Epwarp Burnett Tytor, Oxford, England.

1904. Huco DE Vrizs, Amsterdam, Netherlands.

1907. James WarpD, Cambridge, England.

1909. AuGcusT WEISSMANN, Freiburg, Germany.

1904. WiLHELM WuwnpT, Leipzig, Germany.

CORRESPONDING MEMBERS

31 DECEMBER, 1912.

1883. CHARLES ConraD ABport, Trenton, N. J.

1891. Jost G. AcuitERa, Mexico City, Mexico.

1890. Witi1am De Wirr ALEXANDER, Honolulu, Hawaii.

1899. CO. W. AnprEws, London, England.

1876. JoHN Howarp ApPLeton, Providence, R. I.

1899. J. G. Baker, Kew, England.

1898. Isaac BactEy Batrour, Edinburgh, Scotland.

1878. ALEXANDER GRAHAM BELL, Washington, D. C.

1867. Hpwarp L. BrertHoup, Golden, Colo.

1897. Hersert Bouton, Bristol, England.

1899. G. A. BouLencrr, London, England.

1874. T. 8. Branpscee, Berkeley, Calif.

1884. JoHN C. Branner, Stanford University, Calif.

1894. BonustAy Brauner, Prague, Bohemia.

1874. Wrii1am Brewster, Cambridge, Mass.

1898. T. C. CHAMBERLIN, Chicago, II].

1876. FRANK WIGGLESWORTH CLARKE, Washington, D. C.

1891. L. Cuero, Ekaterinburg, Russia.

1877. THEoporEe B. Comstock, Los Angeles, Calif.

ELECTED.

1868.

1876.

1880.

LSI.

1895.

SiC9.

1870.

1885.

1898.

1894.

1899.

1890.

1899.

1876.

1880.

1869.

1879.

1885.

1899.

1879.

1870.

1858.

1865.

1888.

1868.

1883.

1869.

1898.

1882.

1867.

1900.

1890.

1896.

1875.

1899.

1876.

1888.

1876.

MEMBERSHIP

M. C. Cooxs, London, England.

H. B. CoRNWALL, Princeton, N. J.

CHARLES B. Cory, Boston, Mass.

JOSEPH CRAWForD, Philadelphia, Pa.

Henry P. CusHine, Cleveland, O.

T. Newtson Date, Pittsfield, Mass.

Wiuii1am Hearty Datu, Washington, D. C.

EpwWArD SALispurY Dana, New Haven, Conn.

WitiiamM M. Davis, Cambridge, Mass.

RuTHVEN DeEANzE, Chicago, Il.

CHARLES D&PERET, Lyons, France.

‘ORVILLE A. DrrBy, Rio de Janeiro, Brazil.

Louis Dotto, Brussels, Belgium.

Henry W. Evuiorr, Lakewood, O.

JOHN B. Exxiort, Tulane Univ., La.

Francis EK. ENGELHARDT, Syracuse, N. Y.

Herman Le Roy FarrcHitp, Rochester, N. Y.

FRIEDRICH BERNHARD Firrica, Marburg, Germany.

Lazarus FLetcHER, London, England.

EBERHARD FRAAS, Stuttgart, Germany.

REINHOLD FRITZGARTNER, Tegucigalpa, Honduras.

Grove K. GILBERT, Washington, D. C.

THEODORE NICHOLAS GILL, Washington, D. C.

CuHarLes A. Gorssman, Amherst, Mass.

Frank Austin Goocu, New Haven, Conn.

C. R. GREENLEAF, San Francisco, Calif.

Marquis ANTONIO DE GREGORIO, Palermo, Sicily.

R. J. LecHMErE Guppy, Trinidad, British West Indies.

GrorcE EH. Hatz, Mt. Wilson, Calif.

Baron Ernst von Husse-Wartece, Lucerne, Switzerland.

C. H. HitcxHcock, Honolulu, H. I.

: WiLL1AmM Henry Houmeus, Washington, D. C.

H. D. Hosxoup, Buenos Ayres, Argentine Republic.

J. P. Ippines, Brinklow, Md.

Matvern W. Itzs, Dubuque, Ia.

Orto JAKEL, Greifswald, Germany.

Davip STaRR JoRDAN, Stanford University, Calif.

Grorce A. Kornic, Houghton, Mich.

Baron R. Kuxt, Tokyo, Japan.

JoHN W. LANGLEY, Cleveland, O.

405

406 ANNALS NEW YORK ACADEMY OF SCIENCES

EXLECTED.

1876. S. A. Lartrmore, Rochester, N. Y.

1894. Wuti1am Lipsey, Princeton, N. J.

1899. ARCHIBALD LiversipGE, London, England.

1876. GrorGE Mactoskiz#, Princeton, N. J.

1876. JOHN WILLIAM MALLET, Charlottesville, Va.

1891. CHarLes Rrsore Mann, Chicago, III.

1867. Gzrorce F. MartrHew, St. John, N. B., Canada.

1874. CHARLES JOHNSON Maynarp, West Newton, Mass.

1874. THroporE LuquEER Meap, Oviedo, Fla.

1888. SrrH EH. Mesx, Chicago, Il.

1892. J. DE Menp1zABAaL-TAMBORREL, Mexico City, Mexico.

1874. Ciinton Hart Merriam, Washington, D. C.

1898. MANSFIELD MrrRIAM, South Bethlehem, Pa.

1878. CHARLES SEDGwiIcK Minot, Boston, Mass.

1876. WiLt1am GILBERT MixteR, New Haven, Conn.

1890. RicHarD MoLpENKE, Watchung, N. J.

1895. C. Luoyp Morea, Bristol, England.

1864. Epwarp S. Morss, Salem, Mass.

1898. GrorGE Murray, London, England.

——. Hucren NerTo, Giessen, Germany.

1866. ALFreD Newton, Cambridge, England.

1897. Francis C. NicHotas, New York, N. Y.

1882. Henry ALFRED ALForD NICHOLLS, Dominica, B. W. i,

1880. Epwarp J. Nouan, Philadelphia, Pa.

1879. FREDERICK A. OBER, Hackensack, N. J.

1876. Joun M. Orpway, New Silence, lian

1900. Grorce Howarp Parker, Cambridge, Mass.

1876. StepHEN F. PeckHAm, New York, N. Y.

1877. FREDERICK PRIME, Philadelphia, Pa.

1868. RAPHAEL PuMPELLY, Newport, R. I.

1876. B. ALEx. RANDALL, Philadelphia, Pa.

1876. Ira RemsSEN, Baltimore, Md.

1874. Ropert Ripeway, Washington, D. C.

1886. Witiiam L. Ross, Troy, N. Y.

1876. Samuet P. Sapruer, Philadelphia, Pa.

1899. D. Max Scutossrr, Munich, Germany.

1898. W.B. Scort, Princeton, N. J.

1894. W. T. Sepewicr, Boston, Mass.

1876. ANDREW SHERWOOD, Portland, Ore.

1883. J. Warp SmitH, Newark, N. J.

ELECTED.

1895.

1890.

1896.

1890.

1876.

1885.

1893.

1899.

1877.

1876.

issiral

1900.

1867.

1890.

1898.

1876.

1897.

1874.

1898.

1866.

1899.

1876.

MEMBERSHIP 407

CHARLES H. SmytTu, Jr., Princeton, N. J.

J. SELDEN SPENCER, Tarrytown, N. Y.

Rospert STEARNS, Los Angeles, Calif.

WaLTER LE CONTE STEVENS, Lexington, Va.

Francis H. Storer, Boston, Mass.

Rajah Sourrtnpro Mouun Tacore, Calcutta, India.

J. P. THomson, Brisbane, Queensland, Australia.

R. H. Traqguatr, Colinton, Scotland.

JOHN TROWBRIDGE, Cambridge, Mass.

D. K. Tutti, Philadelphia, Pa.

Henri Van Heurcx, Antwerp, Belgium.

CHARLES R. VAN Hiss, Madison, Wis.

ADDISON EMERY VERRILL, New Haven, Conn.

ANTHONY WAYNE VoebEs, San Diego, Calif.

CHARLES DoOLITTLE WaALcott, Washington, D. C.

Lronarp Watpo, New York, N. Y.

Stuart WELLER, Chicago, II.

I. C. Wuite, Morgantown, W. Va.

Henry SHALER W1LLIAMS, Ithaca, N. Y.

N. H. WInNcHELL, Minneapolis, Minn.

Horatio C. Woop, Philadelphia, Pa.

A. SmitH Woopwakrp, London, England.

ARTHUR WILLIAMS WricHt, New Haven, Conn.

Harry Cricy Yarrow, Washington, D. C.

408

ANNALS NEW YORK ACADEMY OF SCIENCES

ACTIVE MEMBERS

1912

Fellowship is indicated by an asterisk (*) before the name; Life Mem-

bership, by a dagger ({) ;.Patronship, by a section mark (§).

*Abbe, Dr. Cleveland

Abercrombie, David T.

+Adams, Edward D.

Agens, F. G., Sr.

tAlexander, Chas. B.

*Allen, J. A., Ph.D.

Allen, James Lane

*+ Allis, Edward Phelps, Jr., Ph.D.

Ames, Oakes

Anderson, A. A.

Anderson, A. J.C.

*+ Andrews, Roy C.

+ Anthony, R. A.

Arctowski, Dr. Henryk

Arend, Francis J.

+Armstrong, 8. T., M.D.

*Arnold, Felix, M.D.

Ashby, George E.

Astor, John Jacob?

Avery, Samuel P.

+Bailey, James M.

+ Barhydt, Mrs. P. H.

*Barnhart, John Hendley

Barron, George D.

*Baskerville, Prof. Charles

Baugh, Miss M. L.

Beal, William R.

*t Beck, Fanning C. T.

*Beebe, C. William

Beller, A.

+ Bergstresser, Charles M.

*Berkey, Charles P., Ph.D.

Betts, Samuel R.

1 Deceased.

van Beuren, F. T.

*Bickmore, Albert 8., Ph.D.

*Bigelow, Prof. Maurice A., Ph.D.

Bigelow, William 8. |

Bijur, Moses

{Billings, Miss Elizabeth

Billings, Frederick

Bishop, Heber R.

Bishop, Miss Mary C.

Bishop, Samuel H.

*Blake, J. A., M.D. ,

*+Bliss, Prof. Charles B.

+Blumenthal, George

—*Boas, Prof. Franz

Boettger, Henry W.

Bohler, Richard F.

Borup, George?

+Bourn, W. B.

Boyd, James

Brinsmade, Charles Lyman

*Bristol, Prof. Charles L.

Bristol, Jno. I. D.

*SBritton, Prot, Ne ii ene:

*$Brown, Hon. Addison

Brown, Edwin H.

Browne, T. Quincy

*Brownell, Silas B., LL.D.

Bulkley, L. Duncan

Burr, Winthrop

*Bush, Wendell T.

Byrne, Joseph, M.D.

*Byrnes, Miss Esther F., Ph.D.

Camp, Frederick A.

MEMBERSHIP

*Campbell, Prof. William, Ph.D.

*Campbell, Prof. William M.

Canfield, R. A. .

Cannon, J. G.

Carlebach, Walter Maxwell

*SCasey, Col. T. L., U.S. A.

Cassard, William J.

Cassebeer, H. A., Jr.

*+Cattell, Prof. J. McKeen, Ph.D.

§Chapin, Chester W.

*Chapman, Frank M.

+Chaves, José BE.

*Cheesman, Timothy M., M.D.

Childs, Wm., Jr.

Chubb, Percy

Clarkson, Banyer

Cline, M. Hunt

+Clyde, Wm. P.

Cohn, Julius M.

Collier, Robert J.

+Collord, George W.

Combe, Mrs. William -

+Constant, S. Victor

de Coppet, EH. J.

Corning, Christopher, R.

*Crampton, Prof. Henry E., Ph.D.

+Crane, Zenas

Crosby, Maunsell S.

*Curtis, Carlton C.

Curtis, G. Warrington

*Dahlgren, B. E., D.M.D.

Davies, J. Clarence

Davis, Dr. Charles H.

Davis, David T.

~ *+Davis, William T.

*+Dean, Prof. Bashford, Ph.D.

+ Delafield, Maturin L., Jr.

Delano, Warren, Jr. !

Demorest, William C.

Devereux, W. B.

409

De Vinne, Theodore L.

De Witt, William G.

Dickerson, Edward N.

Diefenthaler, C. E.

Dimock, George E.

Dodge, Rev. D. Stuart, D.D.

+ Dodge, Miss Grace H.

*Dodge, Prof. Richard E., A.M.

Doherty, Henry L.

Donald, James M.

*Doremus, Prof. Charles A., Ph.D.

*+ Douglas, James

Douglass, Alfred

Draper, Mrs. M. A. P.

Drummond, Isaac W., M.D.

*Dudley, P. H., Ph.D.

*Dunham, Edward K., M.D.

+Dunn, Gano

+Dunscombe, George Elsworth

*Dutcher, Wm.

*Dwight, Jonathan, Jr., M.D.

Dwight, Mrs. M. E.

*Harle, R. B.

*Hastman, Prof. Charles R.

*+Hilliott, Prof. A. H., Ph.D. :

Emmet, C. Temple

Eno, William Phelps

Estabrook, A. F.

EKvarts, Allen W.

*Hyerman, John

Fairchild, Charles 8.

Fargo, James C.

Farmer, Alexander S.

*Farrand, Prof. Livingston, M.D.

Farrington, Wm. H.

Fearing, D. B.

Ferguson, Mrs. Juliana Armour

§ Field, C. de Peyster

Field, William B. Osgood

*Finley, Pres. John H.

*Fishberg, Maurice, M.D.

410 ANNALS NEW YORK ACADEMY OF SCIENCES

‘Follett, Richard E.

Foot, James D.

+ Ford, James B.

Fordyce, John A.

de Forest, Robert W.

Friedrick, J. J.

Frissell, A. 8.

Fuller, Charles D.

*Gager, C. Stuart, Ph.D.

Gallatin, F.

Gardner, Clarence Roe

Gibson, R. W.

*Gies, Prof. William J.

*Girty, George H., Ph.D.

Godkin, Lawrence

Goodridge, Frederick G.

Goodwin, Albert C.1

§Gould, Edwin

§Gould, George J.

*+Grabau, Prof. Amadeus W.

*Gratacap, Louis P.

Green, James W.

Greenhut, Benedict J.

*Gregory, W. K., Ph.D.

+Grinnell, G. B.

Griscom, C. A., Jr.

Guernsey, H. W.

Guggenheim, William

Guinzburg, A. M.

von Hagen, Hugo

Haines, John P.

Halls, William, Jr.

Hammond, James B.

Hardon, Mrs. H. W.

+Harrah, Chas. J.

+ Harriman, Mrs. E. H.

Hasslacher, Jacob

Haupt, Louis, M.D.

Havemeyer, J. C.

Havemeyer, William F.

1 Deceased.

Healy, J. R.

*Hering, Prof. Daniel W.

Hewlett, Walter J.

*Hill, Robert T.

Hirsch, Charles 8S.

*Hitchcock, Miss F. R. M., Ph.D.

Hochschild, Berthold

Hollenback, Miss Amelia B.

*Hollick, Arthur, Ph.D.

+ Holt, Henry

+ Hopkins, George B.

*Hornaday, Wiliam T., Se.D.

Hotchkiss, Henry D.

*t Hovey, Edmund Otis, Ph.D.

*Howe, Marshall A., Ph.D.

+ Hoyt, A. W.

+Hoyt, Theodore R.

+Hubbard, Thomas H.

Hubbard, Walter C.

Humphreys, Edwin W.

_ Humphreys, Frederic H.

+ Huntington, Archer M.

*Hussakof, Louis, Ph.D.

Hustace, Francis

+ Hutter, Karl

+Hyde, B. Talbot B.

Hyde, E. Francis

+ Hyde, Frederic E., M.D.

Hyde, Henry St. John

*Hyde, Jesse E.

tIles, George

*Trving, Prof. John D.

von Isakovics, Alois

Iselin, Mrs. William E.

+Jackson, V. H.

*Jacobi, Abram, M.D.

James, F. Wilton

tJarvie, James N.

Jennings, Robert H.

*Johnson, Prof. D. W., Ph.D.

MEMBERSHIP

+ Johnston, J. Herbert

Jones, Dwight A.

*S Julien, Alexis A., Ph.D.

Kahn, Otto H.

Kautz-Hulenburg, Miss P. R.

*tKemp, Prof. James F., Sc.D.

+ Keppler, Rudolph

+ Kessler, George A.

Kinney, Morris

Kohlman, Charles

*t Kunz, George F., M.A., Ph.D.

+Lamb, Osborn R.

Landon, Francis G.

Lang, Herbert

Langdon, Woodbury G.

Langeloth, J.

*Langmann, Gustav, M.D.

Lawrence, Amos E.

Lawrence, John B.

+Lawton, James M.

*Ledoux, Albert R., Ph.D.

*Lee, Prof. Frederic 8., Ph.D.

*$Levison, Wallace Goold

Levy, Emanuel

Lichtenstein, M.

Lichtenstein, Paul

lhieb, J. W., Jr.

Lindbo, J. A.

+ Loeb, James

*Loeb, Prof. Morris, Ph.D.?

tLow, Hon. Seth, LL.D.

*Lowie, Robert H., Ph.D.

*Lucas, F. A., D. Se.

*Lusk, Prof. Graham, M.D.

Lydig, Philip M.

Lyman, Frank

_ Lyon, Ralph

McCarthy, J. M.

*tMcMillin, Emerson

McNeil, Charles R.

1 Deceased.

MacArthur, Arthur F.

Macy, Miss Mary Sutton, M.D.

tMacy, V. Everit

Mager, F. Robert

Mann, W. D.

Mansfield, Prof. William

Marble, Manton

Marcou, John B.1

Marling, Alfred H.

t Marshall, Louis

Marston, H.S8.

+ Martin, Bradley

*tMartin, Prof. Daniel 8S.

*Martin, T. Commerford

*+Matthew, W. D., Ph.D.

Maxwell, Francis T.

§ Mead, Walter H.

Mellen, C.S.

*Meltzer, 8. J., M.D.

Merrill, Frederick J. H., Ph.D.

Metz, Herman A.

~ Milburn, J. G. .

Miller, George N., M.D.

*+ Miner, Roy Waldo

Mitchell, Arthur M.

Monae-Lesser, A., M.D.

Morgan, J. Pierpont

*Morgan, Prof. Thomas H.

Morgan, William Fellowes

Morris, Lewis R., M.D.

Munn, John P.

+Nash, Nathaniel C.

+ Nesbit, Abram G.

Notman, George

Oakes, Francis J.

Ochs, Adolph S.

Oettinger, P. J., M.D.

*+ Ogilvie, Miss Ida H., Ph.D.

tOlcott, E. E.

Olmsted, Mrs. Charles T.

412 ANNALS NEW YORK ACADEMY OF SCIENCES

Oppenheimer, Henry S.

*+ Osborn, Prof. H. F., Se.D., LL.D.

Osborn, William C.

+Osborn, Mrs. William C.

*Osburn, Raymond C., Ph.D.

+Owen, Miss Juliette A.

= Pacmis A) beeehaD:

Paddock, Eugene H.*

+ Parish, Henry

Parsons, C. W.

*Parsons, John E.

+Patten, John

Paul, John J.

*Pedersen, Prof. F. M., Ph.D.

*tPellew, Prof. C. E., Ph.D.

Pennington, William *

+ Perkins, William H.

Perry, Charles J.

*Peterson, Frederick, M.D.

Pettigrew, David L.

Pfizer, Charles, Jr.

Philipp, P. Bernard

Phoenix, Lloyd

Pierce, Henry Clay

Plant, Albert

Planten, John R.*

Polk, Dr. W. M.

*Pollard, Charles L., Ph.D.

*Poor, Prof. Charles L.

* Porter, Eugene H.

Post, Abram S.

*Post, C. A.

*Post, George B.

Preston, Veryl

*Prince, Prof. John Dyneley

+Pyne, M. Taylor

*+Ricketts, Prof. P. de P., Ph.D.

Riederer, Ludwig

Robert, Samuel

Roberts, C. H.

1 Deceased.

+ Roebling, John A.

Rogers, E. L.

Rosenbaum, Selig

Rossbach, Jacob ~

tde Rubio, H. A. C.

*tRusby, Prof. Henry H., M.D.

Russ, Edward +

Sachs, Paul J.

Sage, Dean

Sage, John H.

+Schermerhorn, F. A.

Schiff, Jacob H.

Scholle, A. H.

Schoney, Dr. L.

+Schott, Charles M., Jr.

Scott, George S.?

Scoville, Robert

Seaman, Dr. Louis L.

Seitz, Carl E.

Seligman, Jefferson

_ Sexton, Laurence E.

Shaw, Mrs. John C.

Shepard, C. Sidney

§Shepard, Mrs. Finley J.

*Sherwood, George H.

Shillaber, William

Shultz, Charles 8.

*Sickels, Ivin, M.D.

Sleight, Chas. E.

Sloan, Benson B.

Smith, Adelbert J.

*Smith, Ernest E., M.D., Ph.D.

Smith, Frank Morse

Snow, Elbridge G.

*Southwick, Edmund B., Ph.D.

Squibb, Edward H., M.D.

Starr, Louis Morris

*Starr, Prof. M. Allen

*+Stefansson, V.

Steinbrugge, Edward, Jr.

MEMBERSHIP 413

+Stetson, F. L.

*Stevens, George T., M.D.

*+Stevenson, Prof. John J., LL.D.

Stokes, James

Stokes, J. G. Phelps

+Stone, Miss Ellen J.

Straus, Isidor +

Strauss, Charles

Strauss, Frederick

+Streat, James

Sturgis, Mrs. Elizabeth M.

Taggart, Rush

*t+Tatlock, John, Jr.

Taylor, George

Taylor, W. A.

Taylor, William H.

+Terry, James?

Tesla, Nikola

*Thatcher, Edward J., Jr.

Thaw, A. Blair

Thaw, Benjamin

Thompson, Mrs. Frederick F.

Thompson, Lewis S.

+Thompson, Robert M.

*Thompson, Prof. W. Gilman

Thompson, Walter

*Thorndike, Prof. Edward L.

Thorne, Samuel

*Tower, R. W., Ph.D.

*Townsend, Charles H., Sc.D.

Tows, C. D.

*Trowbridge, Prof. C. C.

Tuckerman, Alfred, Ph.D.

Tuttle, Mrs. B. B.

1 Deceased.

Ullmann, E. 8.

+ Vail, Theo. N.

Vanderpoel, Mrs. J. A.

+ Van Slyck, George W.

+Van Wyck, Robert A.

Vreeland, Frederick K.

Walker, William I.

*tWaller, Prof. Elwyn, Ph.D.

Warburg, F. N.

Warburg, Paul M.

Ward, Artemas

+ Ward, Charles Willis

Ward, John Gilbert

Waterbury, J. I.

Watson, John J., Jr.

+ Weir, Col. John?

*Wells, F. Lyman

Williams, R. H.

Wills, Charles T.

*Wilson, Prof. E. B., Ph.D., LL.D.

Wilson, J. H.

Wilson, Miss M. B., M.D.

*Winslow, Prof. Charles-H. A.

*Wissler, Clark, Ph.D.

Woerishoffer, Mrs. Anna

Wood, Mrs. Cynthia A.

Wood, William C.

*Woodbridge, Prof. F. J. E.

*Woodhull, Prof. John F., Ph.D.

*Woodman, Prof. J. Edmund

*Woodward, Prof. R.S.

*Woodworth, Prof. R. 8S.

Younglove, John, M.D.

Zabriskie, George

414 . ANNALS NEW YORK ACADEMY OF SCIENCES

ASSOCIATE MEMBERS

Billingsley, Paul — Kellicott, W. E., Ph.D.

Brown, Harold Chapman, Ph.D. Kirk, Charles T.

Brown, T. C. McGregor, James Howard

Byrne, Joseph P. Montague, W. P., Ph.D.

Fenner, Clarence N., Ph.D. Mook, Charles

Fettke, Chas. R. Moon, Miss Evangeline

Gordon, Clarence E. Northup, Dwight

Jalplena, Jt3 Ins, 1e)n.|D) O’Connell, Miss Marjorie

Haseman, J. D. Rogers, G. Sherburne

Hunter, George W. Stevenson, A. E:

Johnson, Julius M. Wood, Miss Elvira

NON-RESIDENT MEMBERS

*Berry, Edward W. *Lloyd, Prof. F. EH.

Buchner, Edward F. *Mayer, Dr. A. G.

*Bumpus, H. C. Meyer, Adolph

Burnett, Douglass Petrunkevitch, Alexander, Ph.D.

*Davis, William H. 2 PratiseD ies) emele

English, George L. *Ries, Prof. H.

Finlay, Prof. G. I. Reuter, L. H.

Frankland, Frederick W. *Sumner, Dr. F. B.

Hoffman, 8. V. *van Ingen, Prof. G.

Kendig, Amos B. “Wheeler, Wm. Morton

GENERAL INDEX TO VOLUME XXII

Names of Authors and other Persons in Heavy-face Type

Titles of Papers in SMALL CAPS

Active Members, Plection of, 339, 344, 355,

380 :

Active Members, List of, 408

Adams, —, Reference to, 50

Adams, Frank D., Honorary Member, 383

AERIAL TRANSMISSION OF DISEASE, THE,

C. V. Chapin [Abstract], 351, 352

Agassiz, —, Reference to, 86

AGE OF WALKING AND TALKING IN RELATION

TO GENERAL PRACTICE, THE, C. D.

Mead [Abstract], 359, 363 °

ALASKA, GEOLOGY AND MINERAL RESOURCES

or, Alfred H. Brooks [Title], 371

Allen, —, Reference to, 50

Allorisma costatum, 4

gilberti, 5

(Pleurophorella ?) reflerum, 3

reflexum, 3

ALTERATIONS IN THE SNAKE RIVER BASALTS,

Charles T. Kirk [Title], 356

Alula, 4

geinitzi, 3

gen. nov., 3

gilberti, 2, 3, 5

gilberti White?, 5

2 lanceolata, 3

squamulifera, 2, 3, 4, 5

AMALIA FARM METEORITE, THE, Edmund Otis

Hovey [Abstract], 345

Ameginho, —, Reference to, 101

Andrews, Roy C., AN EXPLORATION OF

NORTHEASTERN KoreEA [Title], 375

ANNUAL MEETING, MINUTES OF THE, Edmund

Otis Hovey, 383

Ants of South America, 82

Aquatic migration in South America, 42

Arber, —, Reference to, 84

Archean rocks of South America, 17, 18

Archenhold, F. S., ASTRONOMY, EDUCATION

AND CULTURE [Title], 353

Arctowski, Henry, Active Member, 339

Arges marmoratus from the Republic of

‘ Colombia, 327-333

Arnold, Felix, Fellow, 383

Associate Members, Blection of, 3389, 355,

380

Associate Members, List of, 414

Asst, Jappa, Reference to, 11

Astor, John Jacob, Death of, 370

ASTRONOMY, EDUCATION AND CULTURE, F. S.

Archenhold [Title], 353

ATTEMPT TO MEASURE MENTAL WoRK AS A

PsycHo-DyNAMIC PROCESS, THE, Ray-

mond Dodge [Title], 380

AUDITORY AND VISUAL Memory, A. E. Chris-

lip [Abstract], 346, 347

BACTERIA IN City Dust, THH NUMBER AND

KINDS oF, C.-E. A. Winslow and I. S.

Kligler [Abstract], 351, 352

Balta, José, Reference to, 226 :

Barnard, Edward E., THrE PLANET Mars

[Title], 339

Baskerville, Charles, TUNGSTEN [Abstract],

373

Bassler, —, Reference to, 160.

Bateson, —, References to, 70, 114, 116, 119,

13311

BEDFORD SHALE OF OHIO, GEOLOGICAL AGE

OF THE, George H. Girty, 295-319;

_ [Title], 374

Berkey, Charles P., IS THrRE FAULT CoN-

TROL OF THE HUDSON RIVER COURSE?

[Abstract], 350, 351

SECTION OF GEOLOGY .AND MINERALOGY,

350, 355, 371, 374, 380

Boas, Dr. RADOSAVLJEVICH’S CRITIQUE OF

PROFESSOR, Robert H. Lowie [Ab-

stract], 354

Boas, Franz, A YEAR IN Mexico [Title], 376

Boettger, —, Reference to, 29

Booth, Garret and Blair, Reference to, 336

Borelli, —, cited, 268

Borup, George, Death of, 370

Bowles, James F. B., SANITATION OF THE

PANAMA CANAL [Title], 371

Branner, J. C., cited, 30, 84, 91

References to, 10, 17, 31, 42, 86

Brazilian Coast, Topography of the, 25

Brogger, W. C., cited, 136, 137, 139, 142

References to, 137, 138, 139

Brooks, Alfred H., GmOLOGY AND MINERAL

RESOURCES oF ALASKA [Title], 371

Broom, , References to, 96, 97, 99

Brown, —, Reference to, 115

Brown, Barrington, Reference to, 29

Burke and Pinckney, cited, 181

Burroughs, W. G., cited, 296

(415)

416

BUSINESS MEETINGS, MINUTES oF, Edmund

Otis Hovey, 339, 344, 350, 355, 370,

373, 376, 380

Butters, Roy M., References to, 1, 2

Butts, —, cited, 298, 309

References to, 307, 308, 312

By-Laws of the New York Academy of Sci-

ences, 396

Calandruccio, —, Reference to, 321

Callograptus grabaui sp. noy., 142

Campbell, William, Somn NoTES ON IRON AND

STEEL [Title], 342

Carboniferous fossils of South America, 18

Carrel, Alexis, RESULTS OF THE SUTURE OF

BLOOD VESSELS AND THE. TRANS-

PLANTATION OF ORGANS [Title], 378

CAUSE OF THE TIDES, THR, Charles Lane

Poor [Abstract], 378, 379

CEMENT, METAMORPHISM OF

Albert B. Pacini,

340

CHANGES IN THE BEHAVIOR OF THH HEL DuR-

ING TRANSFORMATION, Bashford Dean,

321-326

Chapin, C. V., THE ARIAL TRAN SAE SS TON

or DisHAsE [Abstract], 351, 352

CHEMICAL ART, PRODUCTS oF, Louis H.

Friedburg [Abstract], 353

Chemical composition of Portland cement,

164

PORTLAND,

161-224; [Title],

Chrislip, A. E., AUDITORY AND VISUAL MEM-

ory [Abstract], 346, 347

Christman, Erwin S., Reference to, 269

Clarke, —, cited, 298, 309

Clarke, F. W., cited, 217

Climbing catfish from the Republic of Co-

lombia, 327-333

Coastal plains of Peru, 228

Collie, —, Reference to, 143

COLORADO, HASTERN, INVERTEBRATE FOSSILS

FROM, George H. Girty, 1-8

Conrad, —, Reference to, 29

Constitution of the New York Academy of

Sciences, 395

Cope, —, cited, 268, 269

References to, 54, 78

Corresponding Members, List of, 404

CORRESPONDING SECRETARY, REPORT OF THE,

Henry E. Crampton, 385

Cox, Charles F., Death of, 344

Crampton, Henry E., REPORT OF THH CORRE-

SPONDING SECRETARY, 385

Crandall, R., cited, 21

References to, 10, 17, 42

Crustacea of South America, 79

CuBAN MARINE FisuEs, NoTres on, John T.

Nichols [Abstract], 358

Culler, A. J.. RELATION OF INTERFERENCE TO

ADAPTABILITY [Abstract], 359, 363

Cumings, E. R., cited, 144

ANNALS NEW YORK ACADEMY OF SCIENCES

Cunningham, —, Reference to, 321

Cushman, A. S., Reference to, 114

Dall, —, quoted, 30

Darwin, —, Reference to, 226

Davenport, —, cited, 122

References to, 114, 126, 127, 128

Davis, —, Reference to, 230

Davis, Frank, Reference to, 11

Day, Francis, cited, 321

Dean, Bashford, Do DEVELOPING EMBRYOS

GivE REAL CLUES TO LINES OF DE-

scent? [Abstract], 372

ON THE CHANGES IN THE BEHAVIOR OF

THH HEL(Conger malabaricus) DURING

Its TRANSFORMATION, 321-326; [Ab-

stract], 372

References to, 11, 327

Deaths, 370, 374, 376

De Koninck, —, Reference to, 299

Derby, O. A., References to, 10, 17, 21, 30, 43

de Ribeiro, Alipo Miranda, Reference to, 38

Devonian fossils of South America, 18

de Vries, Hugo, EXPERIMENTAL HYOLUTION

[Title], 381

References to, 54, 70

DICTYONEMA-FAUNA OF NAvy ISLAND, NEW

BRUNSWICK, ON THE, F. F. Hahn,

135-160

Dictyonema flabelliforme Hichw. (sp.), 136

flabelliforme Hichw. var. acadica Mat-

thew, 137

Dieckmann, G. P., cited, 209

Dieder, —, Reference to, 99

DIFFERENT-TONES AND CONSONANCH, F.

Krueger [Title], 380

DISTRIBUTION OF FERRIC CHLORIDE BETWEEN

ETHER AND AQUEOUS HYDROCHLORIC

AcID AT 25° C., Albert B. Pacini

[Abstract], 378, 379

DISTRIBUTION OF PETROLEUM DEPOSITS IN

PERU, Vernon F. Marsters [Abstract],

350, 351

Do DEVELOPING EMBRYOS Give REAL CLUES

AS TO LINES oF DuscuentT?, Bashford

Dean [Abstract], 372

Dodge, Raymond, THE ATTEMPT TO MEASURE

MENTAL WORK AS A PSYCHO-DYNAMIC

Process [Title], 380

Doherty, Henry L., Treasurer, 350

REPORT OF THE TREASURER, 387

Doncaster, —, Reference to, 113

D’Orbigny, —, Reference to, 226

Drainage of Overlook Mountain, 264

Drirt PEBBLES, THE FOSSILS AND HORIZON

oF THE, F. S. Hintze [Title], 377

Durham, Miss, Reference to, 113

Earle, R. B., Active Member, 344

Fellow, 383

Reference to, 161

Request for grant for research, 377

GENERAL INDEX TO VOLUME XXII

Hast Andean. Sea, Topography of, 30

Echo Lake, 266 —

EpiTor, RHPORT OF THE, Edmund Otis Hovey,

387

EDUCATION AND CULTURE, ASTRONOMY, F. S.

Archenhold [Title], 353

EEL DURING TRANSFORMATION, CHANGES IN

THE BEHAVIOR OF THE, Bashford

Dean, 321-326; [Abstract], 372

Higenmann, —, cited, 36, 51

References to, 11, 16, 52, 58, 57, 59, 65,

67, 74, TT, 321

Emmons, —, Reference to, 140

Eschwege, —, Reference to, 17

Etheridge, —, References to, 29, 30

EURYPTERIDS, PRESENT OPINIONS ON THH

HABITS OF THE, Marjorie O’Connell

\[Title], 377 :

Evans, —, References to, 18, 22

HXPERIMENT IN THE CATCHING OF PENNIES,

E. S. Reynolds, J. T. Gyger and L. L.

Winslow [Abstract], 359, 365

EXPERIMENTAL HVOLUTION, Hugo de Vries

[Title], 381

EXPLORATION OF NORTHEASTERN Korea, AN,

Roy C. Andrews [Title], 375

Ewald, —, Reference to, 281

Fearnside, W. G., cited, 136, 152, 157

Reference to, 158

Fellows, Election of, 383

Fettke, Charles R., Associate Member, 339

Fick, —, cited, 277

Reference to, 281

Fischer, O., Reference to, 268

Foerste, A. F., cited, 304, 318

Foote Mineral Company, Reference to, 336

Foote, Warren M., Reference to, 335

Forbes, —, References to, 101, 226

FORELANDS OF THE BRAS D’OR LAKES, CAPE

BRETON ISLAND, Nova Scotia, J. E.

Woodman [Abstract], 350, 351

FOSSILS AND HorRIZON OF THE DRIFT PEB-

BLES, THE, F. S. Hintze [Title], 377.

Frech, F., cited, 147

Friedburg, Louis H., PrRoDUCTS oF CHEMICAL

Art [Abstract], 353

Gabb, —, Reference to, 29

Gaudry, —, cited, 269

Reference to, 100

GEM-BEARING PEGMATITES OF LOWER CALI-

FORNIA, THE, George F. Kunz [Title],

371 ;

GEOGRAPHICAL DISTRIBUTION IN SOUTH

AMERICA, John D. Haseman, 9-112;

[Title], 346

GEOLOGICAL AGE OF THH BEDFORD SHALES

OF OHIO, George H. Girty, 295-319;

[Title], 374

417

GEOLOGY AND MINERAL RESOURCES OF

ALASKA, Alfred H. Brooks [Title], 371

Geology and topography of South America,

17-49 ;

Gidley, J. W., cited, 286

Girty, George H., GEOLOGICAL AGE OF THE

BEDFORD SHALH OF OHIO, 295-319;

[Title], 374

ON SOME INVERTEBRATE FOSSILS FROM

THN LYKINS FORMATION OF EASTERN

CoLorRADO, 1-8; [Title], 345

Glaciation of Overlook Mountain, 262

Glenn, —, cited, 298, 309

GoAT ISLAND AT NIAGARA GLEN, WAS THERE

A FormMmr, A. W. Grabau [Abstract],

377, 378

Goddard, Henry H., TH HEREDITY OF MEN-

TAL TRAITS [Abstract], 346, 348

Goddard, Pliny E., NOTES ON THE JICARILLA

APACHE [Abstract], 343

Goldbeck, A. T., cited, 208

Goldfarb, A. J.. THE INFLUENCH OF THE

NERvVoUS SYSTEM UPON GROWTH

[Title], 381

A New METHOD OF FUSING EGGS OF

THH SAMBH SpEcIES [Abstract], 381,

382

Gondwana flora of South America, 83

Goodale, H. D., References to, 114, 116

Goodale, H. D., T. H. Morgan and, Sprx-

LINKED INHERITANCE IN POULTRY,

113-133

Goodale, R. C., Reference to, 115

Goodsir, —, Reference to, 273

Goppert, —, cited, 137, 138

Reference to, 152

Gordon, John, Reference to, 11

Grabau, A. W., References to, 11, 136, 138,

148

Was THERE A FORMER GOAT ISLAND AT

NraGarRA GLEN? [Abstract], 377, 378

Grants from research funds, 340, 377

Grassi, —, Reference to, 321

Gregory, —, cited, 55

References to, 11, 82, 99

Gregory, H. E., Reference to, 259 }

Gregory, William K., NoTES ON CERTAIN

PRINCIPLES OF QUADRUPEDAL LOCO-

MOTION AND ON THE MECHANISM OF

THD LIMBS OF HOOFED ANIMALS, 267-

298; Title], 372.

SEcTion oF Brouoey, 340, 346, 351, 358,

871, 375,. 378, 381

Guyer, —, Reference to, 132

Gyger, J. T., E. S. Reynolds and L. L. Win-

slow, EXPERIMENT IN THH CATCHING

OF PENNIES [Abstract], 359, 365

Haacke, —, cited, 17

Reference, 101

Hadley, —, Reference to, 114

418

Hagedoorn, —, Reference to, 114

Hahn, F. F., Associate Member, 355

ON THD DICTYONEMA-FAUNA OF NAVY

ISLAND, Nrw BRUNSWICK, 135-160;

[Title], 372

Hale, George E., Honorary Member, 383

Hall, James, cited, 140

References to, 144, 159, 297, 304, 312

Hardening process of Portland Cement, 166

Hardon, Mrs. Henry W., Active Member, 355

Hartnagel, —, cited, 307

Hartt, —, References to, 17, 25

Haseman, John D., SoMmE FAcTORS OF GEO-

GRAPHICAL DISTRIBUTION IN SoutTH

AMERICA, 9-112; [Title], 346

Hatcher, —, References to, 17, 34

Hauthal, —, Reference to, 17

Haycraft, —, cited, 268, 276, 277, 280, 281

Hedley, —, Reference to, 101

Helmholtz, —, cited, 277

Henke, —, Reference to, 281

HEREDITY OF MENTAL TRAITS, THE, Henry H.

Goddard [Abstract], 346, 348

Herrick, —, cited, 304, 305

Reference to, 306

Hickman, J. E., TH INFLUENCE OF NAR-

COTICS ON PHYSICAL AND MBNTAL

TRAITS OF OFFSPRING [Abstract], 346

Hintze, F. S., TH FOSSILS AND HORIZON OF

THE DRIFT PEBBLES [Title], 377

Holland, W. J., Reference to, 10

Honorary Members, Election of, 383

List of, 403

Hopkinson, T., cited, 140

Horton, B. B., Reference to, 114

Hovey, Edmund Otis, THE AMALIA FARM

MeErTEoRITH [Abstract], 345

THE KINGSTON, N. M., SIDERITE, 335-

337

MINUTES OF THE ANNUAL MEBTING, 383

MINUTES OF BUSINESS MEETINGS, 339,

343, 350, 355, 370, 373, 376, 380

RECORDS OF MEETINGS OF THE NEW

YorRK ACADEMY OF SCIENCES, 339-414

REPORT OF THE EDITOR, 387

REPORT OF THE RECORDING SECRETARY,

385 ~

SECTION OF GEOLOGY AND MINERALOGY,

340, 344, 377

Hupson River Coursn, IS THERE FAULT

CONTROL OF THE, Charles P. Berkey

[Abstract], 350, 351

Hussakof, Louis, Reference to, 11

THE SPAWNING HABITS OF THE SEA

LAMPREY, Petromyzon marinus [Ab-

stract], 358

Huxley, —, Reference to, 101

Hyde, J. E., cited, 296

Fellow, 384

ANNALS NEW YORK ACADEMY OF SCIENCES

‘

ILLUSTRATIONS OF MINERAL ASSOCIATIONS

BY MBANS OF COLOR PLATE AND OTHER

PHOTOGRAPHS OF OPAQUE SPECIMENS,

Wallace Goold Levison [Abstract], 356

INDIVIDUAL DIFFERENCES IN THE INTERESTS '

OF CHILDREN, Gertrude M. Kuper

[Abstract], 359 :

INFLUENCE OF NARCOTICS ON PHYSICAL AND

MENTAL TRAITS OF OFFSPRING, TH,

J. E. Hickman [Abstract], 346

INFLUENCE OF THE NERVOUS SYSTEM UPON

GROWTH, THE, A. J. Goldfarb [Title],

381

INVERTEBRATE FOSSILS FROM HASTERN COLO-

RADO, George H. Girty, 1-8

Invertebrates of South America, 79

Iron, SOME NOTES ON, William Campbell

[Abstract], 343

Is THERD FAULT CONTROL OF THE HUDSON

RIvEK CourSsn?, Charles P. Berkey

[Abstract], 250, 351

Ives, P. P., Reference to, 115

Janda, F., cited, 183

Jay, James E., Reference to, 161

JICARILLA APACHE, NOTES ON THE, Pliny E.

Goddard [Abstract], 343

Johannsen, —, Reference to, 54

Johnson, D..W., Active Member, 380

Fellow, 383 \

THE WESTWARD TRIP OF THE TRANS-

CONTINENTAL WXCURSION OF THB

AMERICAN GEOGRAPHICAL SOCIETY

[Title], 374

Johnson, R. D. O., NoTES ON THH HABITS

OF <A CLIMBING CATFISH (Arges

marmoratus) FROM THE RPPUBLIC OF

CoLomMBIA, 327-333; [Abstract], 372

Jonson, Ernst, Reference to, 161

Jordan, —, References to, 75, 77

Julien, A. A., cited, 262

Katzer, —, References to, 17, 18

, Kemp, James F., References to, 259, 264

King, F. H., cited, 207

Kineston, N. M., SipreritE, THE, Edmund

Otis Hovey, 335-337

T. H., PRACTICE IN THE CASH OF

CHILDREN OF ScHooL AGB [Abstract],

359, 362

Kirk, Charles T., ALTERATIONS IN THD SNAKE

RIvER BASALTS [Title], 356

Kjerulf, Th., References to, 137, 189

Kligler, I. S., C.-E. A. Winslow and, THE

NUMBER AND KINDS OF BACTERIA IN -

Ciry Dust [Abstract], 351, 352

Knorz, —, Reference to, 281

Korpra, AN HXPLORATION OF NORTHEASTERN,

Roy C. Andrews [Title], 375

Krone, Ricardo, References to, 11, 25

Kirby,

: GENERAL INDEX TO VOLUME XXII

Krueger, F., DiIrrrrENT-TONES AND CONSO-

NANCE [Title], 380 (

Kunz, George F., THE GEM-BEARING PEGMA-

TITES OF LOWER CALIFORNIA [Title],

371

Kuper, Gertrude M., INDIVIDUAL DIFFERENCES

IN THE INTERESTS OF CHILDREN [Ab-

stract], 359

Lapworth, Ch., cited, 140

Reference to, 148

Le Chatelier, H., cited, 180

Lesley, Spackmann and, cited, 190

Levison, Wallace Goold, ILLUSTRATIONS OF

MINERAL ASSOCIATIONS BY MEANS OF

CoLorR PLATH AND OTHER PHOTO-

GRAPHS OF OPAQUH SPECIMENS [Ab-

stract], 356

LIBRARIAN, REPORT OF THE, Ralph W. Tower,

386

Tingulella nicholsoni (7), 142

Lisboa, —, Reference to, 91

Loeb, Morris, Death of, 376

Lowie, Robert H., Dr. RADOSAVLJEVICH’S

CRITIQUE OF PROFESSOR Boas [Ab-

stract], 354

Lucas, Frederic A.. WHALING IN THE OLDEN

TIMpB [Title], 341

Luciani, —, cited, 273, 279

Lydekker, —, Reference to, 98

LYKINS FORMATION, ON SOME FOSSILS OF

THE, George H. Girty [Title], 345

Lyon, D. O., THE OPTIMAL DISTRIBUTION OF

TIME AND THE RELATION OF LENGTH

OF MATERIAL TO TIME TAKEN FOR

LEARNING [Abstract], 359, 368

MacDougal, —, Reference to, 70

McGregor, —, References to, 94, 95

McKenna, C. F., cited, 186

MAMMALS, PHYLOGENY AND ONTOGENY OF

THH HorRNS or, H. F. Osborn [Ab-

stract], 341

Mammals of South America, 98

Marey, —, cited, 268, 276, 277

Reference to, 273

Mars, THE PLANET, .Edward EH. Barnard

[Title], 339

Marsters, Vernon F., DISTRIBUTION OF PE-

TROLEUM DmrPoOSITS IN PERU [Ab-

stract], 350, 351

THH PHYSIOGRAPHY OF THE PERUVIAN

ANDES, WitH NoTEes ON HARLY MIN-

ING IN PpruU, 225-258; [Abstract],

345

Matthew, —, References to, 16, 17, 34, 51,

98, 99, 100, 101

Matthew, G. F., cited, 136, 137, 138, 139,

140, 141, 142, 152

References to, 138, 148, 149, 153, 158

419

Matthew, W. D., cited, 270

References to, 135, 137, 269

Maynard, E., cited, 181

Mead, C. D., THmr AGE OF WALKING AND

TALKING IN RELATION TO GENERAL

Practricn [Abstract], 359, 363

Meade, Richard K., cited, 166, 173

Meek, —, Reference to, 304

Membership of the New York Academy of

Sciences, 403-414 :

Memory, AUDITORY AND VISUAL,

Chrislip [Abstract], 346, 347

MENTAL TRAITS, THE HbrREDITY or, Henry

H. Goddard [Abstract], 346, 348

METAMORPHISM. OF PORTLAND CEMENT,

Albert B. Pacini, 161-224; [Title],

340, 356

Metchnikof, Iliya, Honorary Member, 383

METEORITE, TH AMALIA FARM, Edmund

Otis Hovey [Abstract], 345

Mexico, A YHAR IN, Franz Boas [Title], 376

Michaelis, W. A., Sr., and W. A., Jr., cited,

A. &E.

174

Miner, Roy W., TypicAL MARINE INVERTE-

BRATH ASSOCIATION FROM WOODS

HoLE To Casco Bay [Abstract], 375

Mineral constitution of Portland cement, 165

Mining in Peru, Notes on Harly, 254

Mobers, J. Chr., cited, 136, 142

References to, 189, 152, 154

Mollusea of South America, 79

Monobolina refulgens Matthew, 142

Montgomery, Charles M., Reference to, 161

Moodie, —, cited, 95

Moraines of Overlook Mountain, 263

Moreira, Carlos, Reference to, 11

Morgan, T. H., and H. D. Goodale, Smx-

LINKED INHERITANCH IN POULTRY,

113-138 ; [Title], 358

Morse, W. C., cited, 318

Murchisonia buttersi, 2, 8

buttersi sp. nov., 6

lasallensis, 7

terebra, 7

Murray, —, Reference to, 24

Murray, Sir John, Honorary Member, 383

Reference to, 101

Museum or Living Bacteria, A, C.-H. A.

Winslow [Abstract], 381

Muybridge, —, cited, 268

Myalina cuneifornis, 5, 6

perattenuata, 2, 6

perattenuata Meek and Hayden, 6

wyomingensis, 2, 5, 6

wyomingensis Lea, 5

Myers, G. C., Sex DIFFERENCES IN JNCIDEN-

TAL Mmmory [Abstract], 359, 363

Nature of Portland cement, 163

Navy ISLAND, NEw BRUNSWICK, ON THE

DICTYONEMA-FAUNA OF, F. F. Hahn,

135-160 i

420

Neumayer, —, Reference to, 15

Newberry, —, cited, 296

References to, 305, 306

New Mrrnop or FUSING HGGs or THE SAME

Sprecinus, A, A. J. Goldfarb [Abstract],

381, 382

Nichols, —, Reference to, 80

Nichols, John T., NovTES ON CUBAN MARINE

FisHes [Abstract], 358

NITROGEN, RECENT DISCOVERIES CONCERNING

A CHEMICALLY ACTIVE MODIFICATION

or, C. C. Trowbridge [Abstract], 343

Non-Resident Members, List of, 414

Nott ON THE HABITS OF THE CLIMBING

CATFISH (Arges marmoratus) FROM

THE UNITED STATES OF COLOMBIA,

R. D. O. Johnson [Abstract], 372

NOTES ON CERTAIN PRINCIPLES OF QUADRU-

PEDAL LOCOMOTION AND ON ‘THE

MECHANISM OF THR LIMBS’ OF

HOooreD ANIMALS, William K. Gregory

[Title], 372

NOTES ON CUBAN MARINE FisHES, John T.

Nichols [Abstract], 358

NOTES ON THE HABITS OF A CLIMBING CAT-

FISH (Arges marmoratus) FROM THR

REPUBLIC OF CoLoMBIA, R. D. O.

Johnson, 327-333

NOTES ON THH JICARILLA APACHE, Pliny E.

Goddard [Abstract], 343

NOTES ON THE STRUCTURE AND GLACIATION

OF OvyvERLOOK MOUNTAIN, Neil E.

Stevens, 259-266; [Title], 371

NUMBER AND KINDS OF BACTERIA IN CITY

Dust, THE, C.-E. A. Winslow and I. 8S.

Kligler [Abstract], 351, 352

O’Connell, Marjorie, Associate Member, 380

PRESENT OPINIONS ON THE HABITS OF

THE EHURYPTERIDS [Title], 377

ON THE CHANGES IN THE BEHAVIOR OF THD

Eri (Conger malabaricus) DuRING

Its TRANSFORMATION, Bashford Dean

[Abstract], 372

ON THE DICTYONEMA-FAUNA OF Navy Is-

LAND, New BRUNSWICK, F. F. Hahn,

135-160; [Title], 372

On SoMrE FOSSILS oF THE LYKINS FORMA-

TION, George H. Girty [Title], 345

On Somer INVERTEBRATE FOSSILS FROM THE

LYKINS FORMATION OF HASTERN COLO-

RADO, George H. Girty, 1-8

OPTIMAL DISTRIBUTION Or TIME AND THE

RELATION OF LENGTH OF MATERIAL

TO TIME TAKEN FOR LEARNING, D. O.

Lyon [Abstract], 359, 368

Organization of the New York Academy of

Sciences, 389

Origin of the South American Fishes, 75-79

Ortmann, A. E., References to, 11, 16, 79, 80,

81, 82, 101

ANNALS NEW YORK ACADEMY OF SCIENCES

Orton, —, Reference to, 29

Osborn, Henry Fairfield, cited, 16, 52, 78, 98,

102, 269, 284

PHYLOGENY AND ONTOGENY OF THH

Horns or MAMMALS [Abstract], 341

References to, 101, 282, 288

SKULL MEASUREMENTS IN MAN AND

THE HOOFED MAMMALS[Abstract], 341

OvERLOOK MOUNTAIN, NOTES ON THD STRUC-

TURE AND GLACIATION or, Neil E.

Stevens, 259-266; [Title], 371

Pacini, Albert B., Active Member, 339

Tub DISTRIBUTION OF FERRIC CHLORIDD

BETWEEN HTHER AND AQUEOUS HyYDRO-

CHLORIC ACID AT 25° C. Posiesare sl

378, 379

Fellow, 383

METAMORPHISM OF PORTLAND. CEMENT,

161-224; [Title], 340, 356

PANAMA CANAL, SANITATION OF THE, James

F. B. Bowles [Title], 371

Parallelodon, 3

Parsons, Fred H., Reference to, 161

Pearl, —, References to, 114, 122, 131

Pedersen, F. M., SECTION OF ASTRONOMY,

PHYSICS AND CHEMISTRY, 342, 353,

373, 375, 378

Pellegrin, —, cited, 61

Reference to, 54

Pentland, —, Reference to, 226

Permian Inland Basin, Topography of lane

Alto and the, 27

Permian reptiles of South America, 94

PERU, DISTRIBUTION OF PETROLEUM DEPOSITS

IN, Vernon F. Marsters [Abstract],

350, 351

PERU, Harty MINING IN, Vernon F. Mar-

sters, 225-258

Peru, A SKETCH OF THE PHYSIOGRAPHY

AND BWarty MINING DEVELOPMENTS

or, Vernon F. Marsters [Abstract],

345

PERUVIAN ANDES, THE PHYSIOGRAPHY OF

THE, Vernon F. Marsters, 225-258

Peterson, P. M., cited, 190

PETROLEUM D&EPOSITS IN PERU, DISTRIBU-

TION OF, Vernon F. Marsters [Ab-

stract], 350, 351

Pettigrew, —, cited, 272, 273

Phillipi, —, Reference to, 17

PHYLOGENY AND ONTOGENY OF THH HORNS

or Mam™Mats, H. F. Osborn [Ab-

stract], 341

PHYSIOGRAPHY OF THE PERUVIAN ANDES;

Wirn Norms oN Harty MINING IN

Prrvu, Tun, Vernon F. Marsters, 225-

258; [Abstract], 345

Pilsbry, cited, 16, 32, 53

Reference to, 32.

Pinckney, Burke and, cited, “181

GENERAL INDEX TO VOLUME XXII

Pissis, —, Reference to, 226

PLANET Mars, THn, Edward E. Barnard

[Title], 339

Plano Alto and the Permian Inland Basin,

Topography of, 27

Pleurophorella, 3

Pleurophori, 3

Pleurophorus, 3

geinitzi, 3

gilberti, 3

lanceolata, 3

papillosa, 3, 4

sp., 2, 6

Poctta, —, Reference to, 160

Poincaré, Jules Henri, Death of, 374

Poor, Charles Lane, THE CAUSE OF THE

Tipes [Abstract], 378, 379

PORTLAND CHMENT, METAMORPHISM oF,

Albert B. Pacini, 161-224; [Title],

340, 356

Poulsen, A., cited, 180

POULTRY, SHPX-LINKED INHERITANCE IN, T. H.

Morgan and H. D. Goodale, 113-133;

[Title], 358

PRACTICH IN THE CASH OF CHILDREN OF

ScHoot Acu, T. H. Kirby [Abstract],

359, 362

PRESENT OPINIONS ON THE HABITS OF THE

EURYPTERIDS, Marjorie O’ Connell

[Title], 377

Price, Charles E., Reference to, 161

PropuctTs OF CHEMICAL ART,

Friedburg [Abstract], 353

Prosser, —, Reference to, 318

PSYCHOLOGY OF THE PHARTHWORM,

Robert M. Yerkes [Title], 380

Punnett, —, Reference to, 114

Louis H.

THE,

Quadrupedal locomotion, 267-294

-RADOSAVLJEVICH’S CRITIQUE OF PROFESSOR

Boas, Robert H. Lowie [Abstract],

354

Raimondi, —, References to, 226, 227, 247,

248

Raynor, —, Reference to, 113

RECENT DISCOVERIES CONCERNING A CHEM-

ICALLY ACTIVH MODIFICATION OF

NITROGEN, C. C. Trowbridge [Ab-

stract], 343

RECORDING SECRETARY, REPORT OF THE,

Edmund Otis Hovey, 385

RECORDS OF MEETINGS OF THE NEW YORK

ACADEMY OF SCIENCES, Hdmund Otis

Hovey, 339-414

Regan, C. Tate, cited, 328

Reid, Harry Fielding, THr SEISMOGRAPH

AND WHAT It THacHEs [Abstract],

381

RELATION OF THH FELDSPARS, LENADS AND

VnoLtites, Henry S. Washington [Ab-

stract], 345

421

RELATION OF INTERFERENCE TO ADAPTABIL-

ity, A. J. Culler [Abstract], 359, 363

Report of the Corresponding Secretary, 385

Editor, 387 ;

Librarian, 386

Recording Secretary, 385

Treasurer, 387

Reptiles of South America, 94

RESULTS OF THE SUTURE OF BLOOD VESSELS

AND THH TRANSPLANTATION OF OR-

GANS, Alexis Carrel [Title], 378

Reynolds, E. S., J. T. Gyger and L. L. Win-

slow, EXPERIMENT IN THE CATCHING

OF PENNIES [Abstract], 359, 365

de Ribeiro, Alipio Miranda, Reference to, 11

Rich, J. L., cited, 264 -

Rio Amazonas, Reversal of the, 33

Roemer, F., cited, 147

Rohland, P., cited, 179, 182

Roth, —, Reference to, 100

Ruedemann, R., cited, 136, 137, 139, 140,

141, 143, 145, 146, 147, 149, 152

References to, 136, 137, 144, 147, 148,

+ 157, 159, 160

Riitimeyer, —, Reference to, 101

Ryder, -—, cited, 268

Salter, —, cited, 138, 142

Reference to, 152

Sanderson, J. Burdon, cited, 282

SANITATION OF THE PANAMA CANAL, James

F. B. Bowles [Title], 371

Sanquinolites, 4

Schafer, E. A., cited, 268

Scheuzer, —, Reference to, 92

Schmaltz, R., cited, 292

Schuchert, Charles, cited, 306

References to, 10, 16, 99, 263, 306

Sclater, —, Reference to, 98

Spa LaAmMprey, Petromyzon marinus,

SPAWNING HABITS OF THE,

Hussakof [Abstract], 358

SECTION OF ANTHROPOLOGY AND PSYCHOLOGY,

F. Lyman Wells, 343, 346, 354, 359,

376, 380

SECTION OF ASTRONOMY, PHYSICS AND

CHEMISTRY, F. M. Pedersen, 342, 353,

373, 375, 378

SECTION oF BroLoGgy, William K. Gregory,

340, 346, 351, 358, 371, 375, 378, 381

SECTION OF GEOLOGY AND MINERALOGY,

Charles P. Berkey, 350, 355, 371, 374,

380

SECTION OF GEOLOGY AND MINERALOGY,

Edmund Otis Hovey, 340, 344, 377

Segerberg, C. O., cited, 136

SHISMOGRAPH AND WHAT IT TEACHES, THE,

Harry Fielding Reid [Abstract], 381

Setting process of Portland cement, 165

Spx DIFFERENCE IN INCIDENTAL MbMOryY,

G. C. Myers [Abstract], 359, 363

THE

Louis

422

SEx-LINKED INHERITANCE IN Pouutry, T. H.

Morgan and H. D. Goodale, 113-133

Sharff, —, Reference to, 101

Siderite from Kingston, N. M., 335-337

Silurian fossils of South America, 18

Simon, Feliciano, Reference to, 11

Sinclair, —, References to, 99, 101

SKETCH OF THN PHYSIOGRAPHY AND HARLY

MINING DEVELOPMENTS oF Prru, A, Ver-

non F. Marsters [Abstract], 345

SKULL MEASUREMENTS IN MAN AND THH

HooreD MAMMALS, H. F. Osborn [Ab-

stract], 341

Sleight, Charles H., Active Member, 344

Smith, A. Erskine, cited, 166

Smock, John C., cited, 266

SNAKE RIvEeR BASALTS, ALTERATIONS IN THH,

Charles T. Kirk [Title], 356

Snetlager, —, Reference to, 11

Some Factors oF GHOGRAPHICAL DISTRIBU-

TION IN SouTH AMERICA, John D.

Haseman, 9-112; [Title], 346

Somn Nores on IRON AND STEEL, William

Campbell [Abstract], 343

South America, Geology and Topography of,

17-49 ;

SoutH AMERICA, SOME FACTORS OF GEO-

LOGICAL DISTRIBUTION IN, John D.

Haseman, 9-112; [Title], 346

South American fishes, Distribution of, 50-61

Spackmann and Lesley, cited, 190

SPAWNING HABITS OF THH SEA LAMPREY,

Petromyzon marinus, TH, Louis

Hussakof [Abstract], 358

Spillman, W. J., References to, 113, 114, 116,

119

Staurograptus dichotomus

apertus Rued., 140

Street, Some Notes on, William Campbell

[Abstract], 343

Stefansson, V., Fellow, 383

Hmmons var.

Steindachner, —, Reference to, 66

Steinmann, G., Reference to, 227

Steinmann, —, References to, 17, 36

Stelzner, —, References to, 22, 31

Stevens, Neil E.. NOTES ON THE STRUCTURD

AND GLACIATION OF OVERLOOK MOUN-

TAIN, 259-266; [Title], 371

Stillman, J. D., cited, 268, 272, 277, 287, 293

Straus, Isidor, Death of, 370

Stream piracy in South America, 38

Structure of Overlook Mountain, 260

Sturtevant, —, Reference to, 114

Suess, —, References to, 15, 17, 22

Surface, —, References to, 114, 122

Tarr, Ralph Stockman, cited, 260

Taylor and Thompson, cited, 173, 180

Thompson, Taylor and, cited, 173, 180

Thompson, G. W., cited, 190

Tiprs, THE CAUSE OF THE, Charles Lane

Poor [Abstract], 378, 379

ANNALS NEW YORK ACADEMY OF SOIENCES

Topographic expression as related to the

geology of the Peruvian Andes, 249

Topographic Provinces of Peru, 227

Tower, —, Reference to, 101

Tower, Ralph W., RerorT or THD LIBRARIAN,

386

Tower, W. I., cited, 49, 50, 54

Reference to, 50

Treasurer, Election of, 350

TREASURER, REPORT OF THE, Henry lL. Doh-

erty, 387 :

Trend lines of South America, 21

Trowbridge, C. C., RrceNT Discoverius Con-

CERNING A CHEMICALLY ACTIVH Mopt-

FICATION OF NITROGEN [Title], 342

Tullberg, S. A., cited, 137, 138

Reference to, 138

TUNGSTEN, Charles Baskerville [Abstract],

373

TYPICAL MARINE INVERTEBRATE ASSOCIATION

FROM WOODS HOLE TO Casco Bay,

Roy W. Miner [Abstract], 375

Ulrich, —, Reference to, 143

Van Hise, C. R., cited, 165, 201

van Ingen, G., Reference to, 135

Var. conferta Linrs. (ms.), 139 —

Var. desmograptoidea var. nov., 139

Var. norwegica Kjerulf., 139

Var. ruedemanni nom. noy., 139

von Huene, —, Reference to, 94, 96

“von Ihering, Rudolfo, References to, 11, 15,

16, 52, 80, 82, 101

Wallace, —, References to, 15, 98

Walther, —, Reference to, 148

WAS THERE A FORMER GOAT ISLAND AT

NIAGARA GLEN ?, A. W. Grabau [Ab-

stract], 377, 378

Washington, Henry S., RELATIONS OF THE

FELDSPARS, LENADS AND YEHOLITES

[Abstract], 345

Watasé, Sho, Honorary Member, 3838

Water upon metamorphism of Portland ce-

ment, Influence of, 168

Weber, Eduard, cited, 280

Weber, Eduard and Wilhelm, cited, 268

Reference to, 273 :

Weir, John, Death of, 370

Weller, —, cited, 313, 316

References to, 314, 315, 318

Wells, F. Lyman, Fellow, 383

SECTION OF ANTHROPOLOGY AND PSY-

CHOLOGY, 343, 346, 354, 359, 376, 380

Westergard, A. H., cited, 136, 137, 138, 139,

141, 146

References to, 137, 148, 152, 153, 155,

158, 159

WESTWARD TRIP OF THE TRANSCONTINENTAL

EXCURSION OF THE AMERICAN GEO-

GRAPHICAL Socipty, THH, D. W. John-

son [Title], 374

GENERAL INDEX TO VOLUME XXII

WEALING IN THE OLDEN TIMB, Frederic A.

Lucas [Title], 341

White, A. H., cited, 171, 208

White, David, References to, 10, 16, 83, 84,

89, 90, 93

White, I. C., cited, 309

Reference to, 83

Wig, R. J., cited, 186

Wiman, Carl, cited, 147

References to, 148, 149, 159, 160

Winslow, C.-E. A., A MusmumM or LiviNnG

Bacteria [Abstract], 381

Winslow, C.-E. A., and I. S. Kligler, THE

NUMBER AND KINDS OF BACTERIA IN

City Dust [Abstract], 351, 352

Winslow, L. L., EH. S. Reynolds and J. T.

Gyger, EXPERIMENT IN THE CATCHING

OF PENNIES [Abstract], 359, 365

423

Woodman, J. E., FORELANDS Or THE Bras

D’OR LAkns, CAPE BrRETON ISLAND,

Nova Scoria [Abstract], 350, 351

Reference to, 161

Woodward, —, cited, 76

References to, 29, 54, 77

Woodworth, —, Reference to, 86

Worthenia, 7, 8

tabulata, 7, 8

Wortman, —, Reference to, 101

Wright, —, Reference to, 115

YEAR IN Mpxico, A, Franz Boaz [Title], 376

Yerkes, Robert M., THE PSYCHOLOGY OF THE

HARTHWORM [Title], 380

Ziegler, Victor, Reference to, 143

Zirkel, Ferdinand, Death of, 374